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t; 






A MONOGRAPH 






ON THE 



FOSSIL LEPADID^E, 



OK, 



PEDUNCULATED CffiRIPEDES OE GREAT BRITAIN. 



BY 



CHARLES DARWIN, F.R.S., F.G.S. 




ZA-ZZSq 









LONDON: 

PRINTED FOR THE PALM)NTOGRAPHICAL SOCIETY. 
1851. 



C. AND J. ADLARD, PRINTERS, BARTHOLOMEW CLOSE. 



PREFACE. 

I have great pleasure in returning my most sincere thanks to various naturalists, 
both for intrusting to me their collections of Fossil Cirripedia, and for allowing me, 
whenever it was advisable, to clear the specimens from their matrix. Although an entire 
stranger to many of the gentlemen to whom I applied, I have in every instance received 
the most courteous acquiescence to my demands. To Mr. Fitch, of Norwich, I here beg- 
to return my thanks, for having allowed me to keep, during several months, his unrivalled 
collection of Cirripedia from the Upper Chalk of Norwich, — the fruit of twenty years' 
labour. Mr. Bowerbank has given me the freest use of his fine collection, rich in 
specimens from the Gault. Mr. Wetherell placed in my hands his beautiful and unique 
specimen of Loricula pulchetta, and other species. Professor Buckman sent me, of his 
own accord, a fine series of the valves of Pollicipes oolitims, the most ancient Cirripede 
as yet known, discovered and named by him. To Messrs. Flower, Searles Wood, 
F. Edwards, Harris, S. Woodward, Tennant, and other gentlemen, I owe the examination 
of several species new to me. Mr. Morris and Professor E. Forbes have, in their usual 
kind manner, supplied me with much valuable information, and with the loan of many 
specimens. To Mr. James de C. Sowerby I must express my thanks for the valuable aid 
rendered to me by the loan of the original specimens figured in the ' Mineral Conchology;' 
and for the pains exhibited in the drawings here published. 

Professor Forchhammer, of Copenhagen, not only placed at my disposal many valuable 
specimens deposited in the Geological Museum of the University, but applied to Professor 
Steenstrup, who, in the most generous manner, sent me the collection in the Zoological 
department, including the highly valuable original specimens of his excellent Memoir on 
the Fossil Cirripedia of Denmark and Scania. Subsequently, Professor Steenstrup sent 
me a second large collection, the fruit of the indefatigable labours of M. Angelin, in 



vi PREFACE. 

Scania : all these northern specimens have been of the greatest use to me in illustrating 
the' British species. Having applied to Professor W. Dunker, of Cassel, for some of the 
species described by various German authors, he not only sent me many specimens out of 
his own collection, but procured from Messrs. Roemer, Koch, and Philippi, other specimens 
of great value ; and to these most distinguished naturalists I beg to return my very 
sincere thanks. Lastly, I may be permitted to state, that I hope very soon to have 
another and more appropriate opportunity of publicly expressing my gratitude to various 
gentlemen, who for many months together have left in my hands their large and valuable 
collections of recent Cirripedia, and who have assisted me in every possible way. I will 
here only state, that it was owing to the suggestion and encouragement of Mr. J. E. Gray, 
of the British Museum, that I was first induced to take up the systematic description of 
the Cirripedia, having originally intended only to study their anatomy. To all the 
foregoing gentlemen, I shall ever feel under the deepest obligations. 



INTRODUCTION. 



The Cirripedia, both recent and fossil, have been much neglected by systematic 
naturalists : the fossil species have, however, been more attended to than the recent. 
Professor Steenstrup has published 1 an excellent monograph on the Danish and Scanian 
Cretaceous species : Mr. J. de Carle Sowerby has given good plates of several British 
valves in the Mineral Conchology ; and F. Roemer 2 has illustrated, by rather indifferent 
figures, though clear descriptions, various German forms. Other less important notices 
have appeared by several authors. As yet, however, no monograph has been produced 
on the whole group. The present volume is confined to the Lepadidce or Pedunculated 
Cirripedia ; and it so happens that the introduction, under the form of notes, of a few 
foreign species (which are necessary to illustrate the British species), serves to render 
this Monograph tolerably complete ; that is, as far as the specimens collected on the 
Continent (judging from published accounts) serve for this end, — for we shall immediately 
see that certain valves are requisite in each genus. 

It is unfortunate how rarely all the valves of the same species have been found 
coembedded ; it is evident that, with the exception of some few species, the membrane 
which held the valves together, decayed very easily, as it does in recent Pedunculated 
Cirripedes. Hence, in the great majority of cases, the several valves have been found 
separate. Hitherto it has been the practice of naturalists to attach specific names 
indifferently to all the valves ; and as in each species there are from three to five or six 
different kinds of valve, there would have been, had not the whole group been much 
neglected, so many names attached to each species. On the other hand, it has occurred 
in several instances, that many valves belonging to quite different species have been 
grouped together under the same name. To avoid these great evils, I have fixed on the 
most characteristic valves, one in each of the two main genera, and taking them as 

1 Naturhistorisk Tidsskrift, af H. Kroyer, 1837 and 1839. 

2 Die Versteinerungen des Norddeutschen Kreidegebirges, 1841. 






2 FOSSIL CIRRIPEDIA. 

typical, have never, except in one instance where several valves were known all to belong 
to the same individual, and in another instance in which a valve was very remarkable, 
attached a specific name to any other one. I have, however, in two cases retained names 
already given to certain other valves, as they presented remarkable characters, and were 
almost certainly distinct. In Scalpellum I have taken the Carina or Keel-valve (i. e. dorsal 
valve of most authors) as typical ; and in Pollicipes, the Scuta (i. e. the inferior lateral 
valves of most authors) : it would have been desirable to have taken the same valve in 
both genera ; but it so happened that the Carina has been much more frequently collected 
than any other valve in Scalpellum, in which genus it is highly characteristic ; whereas 
in Pollicipes, it is apt to present less striking characters than the Scuta, which are, 
moreover, commoner in most collections. In almost all the Lepadidae the Terga (i. e. the 
upper or posterior lateral valves) are not characteristic, and are particularly liable to 
variation. Although only certain valves in each genus thus receive specific names, yet 
from the conditions of embedment, several of the other valves can often be safely 
attributed to the same species. 

Much confusion in nomenclature will, I think, be avoided by the plan here adopted ; 
but the study of Possil Cirripedia must, I fear, owing to the variability of the valves, as 
seen in some fossil species, and as inferred from what so commonly occurs with recent 
species, ever remain difficult. In very many of those recent species, of which large series 
have passed through my hands, several of the valves have varied so much, that had I 
seen only certain specimens from the opposite poles of the series, I should unhesitatingly 
have ranked them as quite distinct species-, on the other hand there are some recent' 
forms — for instance, some species of Zepas, and again Pollicipes cornucopia, and elegans 
of Lesson — which are perfectly distinct, but which it would be hopeless to attempt- 
discriminating when fossilized, without quite perfect specimens. It should be borne in ; 
mind, that the recognition of the Fossil Pedunculated Cirripedes by the whole of their 
valves and peduncle, is identical with recognising a Crustacean by its carapace, without 
the organs of sense, the mouth, the legs, or abdomen : to name a Cirripede by a single 
valve is equivalent to doing this in a Crustacean by a single definite portion of the 
carapace, without the great advantage of its having received the impress of the viscera 
of the included animal's body : knowing this, and yet often having the power to identify ' 
with ease and certainty a Cirripede by one of its valves, or even by a fragment of a 
valve, adds one more to the many known proofs of the exhaustless fertility of Nature in 
the production of diversified yet constant forms. 

I must a]lude to one more unfortunate cause of doubt in the classification of the 
extinct Lepadidae, namely, the difficulty in attributing the separated valves to the two 
main genera of Scalpellum and Pollicipes ; for the chief distinction between these two 
close genera in the recent state, lies in the number of the valves, and this can very rarely 
be ascertained in fossil specimens. At first I determined to follow those authors who 
have united both genera under Pollicipes ; but reflecting that I had twelve recent and 



. INTRODUCTION. 3 

above thirty-seven fossil species, with almost the certainty — as we shall presently see — of 
very many more being discovered, this plan seemed to me too inconvenient to be followed. 
There are six recent species which I intend, in a future work, to include under Scalpellum. 
Four of them have been raised by Dr. Leach and Mr. Gray to the rank of genera • two 
other unnamed species have certainly equal, if not stronger, claims to the same rank ; 
so again the six recent species of Pollicipes have similar claims to be divided into three 
genera, thus making nine genera for the twelve recent species of Scalpellum and 
Pollicipes. In the majority of cases it would be eminently difficult to allocate the fossil 
species in these nine genera ; nevertheless, taking the characters necessarily used for the 
generic divisions of all the other recent Pedunculated Cirripedes, there can be no doubt 
that the formation of the above nine genera might be justified, that is, if we are allowed 
to neglect mere classificatory utility as an element in the decision, and further, if we are 
invariably bound to make as far as possible all genera of exactly the same value. As 
far as utility in classification is concerned, it appears to me clear that the institution of so 
many genera, until many more species are discovered, is highly disadvantageous : with 
respect to making all genera of exactly equal value, this, though eminently desirable, 
appears to me almost hopeless ; I know not how to weigh the value of slight differences 
in different valves ; or whether a difference in the maxillae or mandibles be the more 
important : anyhow, in this particular case, if we raised the six recent species of Scalpellum 
into six genera, they assuredly would not be distinct to an exactly equal degree. Under 
these circumstances I have followed a middle term, and kept Scalpellum and Pollicipes 
distinct, — genera easy to be recognised in a recent state, — which renders the classification 
of the fossil species, though always difficult and liable to many errors, somewhat easier 
than if both genera were united into one, and much easier than if the above nine genera 
were admitted. 



Aptychus. 

Before passing to more general considerations, I must offer a few remarks on the 
genus Aptychus, or Triyonellites, inasmuch as quite lately a distinguished naturalist, 
M. D'Orbigny, 1 has adopted, and with much ingenuity supported, the view that these 
anomalous bodies are Pedunculated Cirripedia. It cannot be denied that the general form 
and lines of growth closely resemble those of the Scuta or lateral inferior valves in Lepas 
or Anatifa : nor can it be denied, from what we know of recent species, that the Terga 
(upper lateral valves) and Carina (dorsal valve), which on M. D'Orbigny's view must be 
considered as absent, are the most likely valves to disappear from abortion. But there 
are points of difference which, as it appears to me, are of far greater importance than the 

1 Cours Elementaire de Palecmtologie, 1849, vol. i, p. 254. 



4 FOSSIL CIRRIPEDIA. 

resemblance in mere outline. The peculiar cancellated structure, which is almost visible on 
the external surface even to the naked eye, is wholly unlike anything known amongst 
Cirripedia ; a thin polished slice of the valves of Lepas and of Aptychus, viewed under a high 
power, are as unlike as anything can well be. 1 In Aptychus the lines of growth are 
conspicuous on the inner or concave surface, and indistinguishable or not plain on the 
outer surface ; whereas in Lepas exactly the reverse holds good. Again, in some 
specimens it appears, that additions are made to the shell on the exterior edge of the 
growing margin, instead of on the inner edge, as in Cirripedia. In Aptychus latus, 
there is a rather deep internal fold along the whole of that margin, through which the 
cirri are supposed to have been protruded, and this is unlike anything which I have met 
with in Cirripedes. In all the species of Aptychus, the two valves are much the most 
frequently, though not invariably, found widely opened, and attached together, either 
exactly or nearly so, by the two margins through which the cirri must have been protruded. 
Now in all true fossil pedunculated Cirripedes, the valves are found either separate, which 
is the commonest case, or when held together, those on the opposite sides almost exactly 
cover each other, for there is nothing in the structure of Cirripedia tending to open the 
valves like the ligament in bivalve shells. How comes it, then, that the specimens of 
Aptychus, even those found within the protected chambers of Ammonites, thus generally 
have their valves widely gaping ? Even if we pass over this difficulty, is it not strange 
that the valves should always have been held together by that margin, which in the recent 
condition is supposed to have been open for a considerable portion of its length, for the 
exsertion of the cirri ; whereas, in not one single instance, as far as I have seen, are the 
two valves held together by the opposite margin, which in the recent state, on the 
idea of Aptychus having been a Cirripede, must have been continuously united by 
membrane. 

There is another argument against Aptychus having been a Cirripede, which will have 
weight, perhaps, with only a few persons : in Pollicipes, the main growth of all the valves 
is downwards ; in Lepas or Anatifa, as well as in most of the allied genera, the main 
growth of the Scuta and of the Carina (*". e. lower lateral, and dorsal, or valves,) is in a 
directly reversed direction, or upwards. Now Pollicipes is the oldest known genus of 
Cirripedes, having been found in the Lower Oolite, whereas hitherto Lepas is not 
certainly known to have been discovered even in the newest Tertiary formation. So again 
within the limits of the genus Scalpellum, I know of only two cretaceous species in which 
the Scuta grow upwards and downwards, and only one case in which the Carina has this 
double direction of growth ; whereas in the recent and one Miocene species, these valves 
usually grow both upwards and downwards. Hence it would appear that there is some 
relation between the age of fossil Lepadidae and the upward or downward direction of 

1 When I had the slices made, I did not know of H. von Meyer's paper on Aptychus, in the 'Acta 
Acad. Caes. Leop. Car.,' vol. xv, Oct. 1829, tab. lviii and lix, fig. 13, in which perfectly accurate sections 
are given of the microscopical structure of Aptychus Icevis. 



INTRODUCTION. 5 

the lines of growth in their valves. Aptychus, according to M. D'Orbigny, existed during the 
Carboniferous system, at a period vastly anterior to the oldest known Pollicipes, yet on 
the idea of its having been a Cirripede, the growth of its valves (Scuta) must have been 
upwards, as in the most recent forms ; and it was allied to Lepas, that genus which, in the 
order of creation, and in the manner of growth, stands at the opposite end of the series 
from Pollicipes. Prom the several reasons now given, it does not appear to me that 
Aptychus, until weightier evidence is adduced, can be safely admitted as a Cirripede. 

Geological History. — No true Sessile Cirripede 1 has hitherto been found in any 
Secondary formation ; considering that at the present time many species are attached to 
oceanic floating objects, that many others live in deep water in congregated masses, that 
their shells are not subject to decay, and that they are not likely to be overlooked when 
fossilized, this seems one of the cases in which negative evidence is of considerable value. 
Mr. Samuel Stutchbury, moreover, (to whom I am deeply indebted for much information, 
and the loan of his beautiful collection of recent species,) has assured me that vast numbers 
of fossil secondary corals have passed through his hands, and that he has carefully looked 
without success for those genera which commonly inhabit living corals. Sessile Cirripedes 
are first found in Eocene deposits, and subsequently, often in abundance, in the later 
Tertiary Formations. These Cirripedes now abound so under every zone, all over the 
world, that the present period will hereafter apparently have as good a claim to be called 
the age of Cirripedes, as the Palaeozoic period has to be called the age of Trilobites. There 
is one apparent exception to the rule that Sessile Cirripedes are not found in Secondary 
formations, for I am enabled to announce that Mr. J. de C. Sowerby has in his collection 
a Verruca (== Clisia, Clytia, Creusia, Ochthosia) from our English chalk : but this genus, 
though hitherto included amongst the Sessile Cirripedes, must, when its whole organisation 
is taken into consideration, be ranked in a distinct family of equal value with the Balanidse 
and Lepadidae, but perhaps more nearly related to the latter than to the Sessile Cirripedes. 
Hence the presence of Verruca in the Chalk is no real exception to the rule that Sessile 
Cirripedes do not occur in Secondary formations ; on the contrary, it harmonises with the 
law, that there is some relation between serial affinities of animals, and their first appearance 
on this earth. 

The oldest known pedunculated Cirripede is a Pollicipes, discovered by Professor 
Buckman in the Stonesfield Slate in the Lower Oolite : two species of the same genus 
have been described by Mr. Morris from the Oxford Clay, in the middle Oolite. I have 

1 Dr. Petzholdt has described and figured (Jahrbuch, 1842, p. 403, tab. x), a Balanus carbonaria 
from the carboniferous system ; but as neither tbe operculum, the structure of the shell, the number of 
the valves, nor their manner of growth, can be made out or are described, the evidence appears quite 
insufficient to admit the existence of this genus at so immensely a remote epoch. Bronn, in the ' Index 
Palaeontologicus,' gives, under Tubicinella, a cretaceous species ; I have unfortunately not been able to 
consult the original work cited. 



6 FOSSIL CIRRIPEDIA. 

not heard of any Cirripede having been as yet discovered in the Upper Oolite, or in the 
Wealden formation. During the deposition of the great Cretaceous System, the Lepadidas 
arrived at their culminant point ; there were then three genera, and at least thirty-two 
species, some occurring in every stage of this system. Besides the thirty-two certainly 
known cretaceous forms, and several other doubtful ones, I believe that very many more 
will yet be discovered ; I infer this from the fact, that in almost every collection lent to 
me for examination, although very small, I have found some new species. I have three 
or four species from the Gault ; from five to eight in the Lower Chalk, and from nine to 
twelve species in the Upper Chalk (not including the Faxoe, Scanian, and Maastricht 
stage) ; and of these nine to twelve species, five have been found by one collector, Mr. Fitch, 
in one locality, namely near Norwich. In Scania M. Angelin has found no less than nine 
or ten species, all belonging to the upper or Maastricht stage of the Chalk. These fossils, 
judging from the habits of recent species of the same genera, were probably attached to 
fixed, or nearly fixed, objects at the bottom of the sea. Now at the present day, of attached 
Pedunculata (reckoning even Crustacea andEchinidge as fixed objects), the whole Mediter- 
ranean and New Zealand can boast each only of three species, in both cases including 
Alepas, which is destitute of calcified valves and therefore not likely to be fossilized; 
Australia has three species ; Madeira has four species, including one with very small and 
imperfectly calcified valves ; the great Phillipine Archipelago, however, has afforded, owing 
to the labours of Mr. Cuming, as many as five species, though including one with horny 
valves, and a Lithotrya which lives embedded on the beach. Therefore since we already 
have nine or ten fossil species from one locality, and from the same stage of the chalk, 
we may admit that the pedunculated Cirripedes arrived during the upper part of the 
Cretaceous system at their culminant point. 

Although, for this family, the number of species were considerable during the Cretaceous 
period, the individuals were mostly rare. I infer this from the small number of specimens 
in all collections ; for instance, Mr. Fitch, who has assiduously collected for twenty years 
in the chalk near Norwich, possesses in his entire collection only nine keel-valves of Scal- 
pellum maximum, and six of S.fossula; he has two Scuta (and with regard to these valves, 
it must be remembered, that each individual had two) of Pollicipes striatus, two of 
P.fallax, and four of P. Angelini. This occasional want of a relation, within the same 
region, between the number of the species in any given genus, and of the individuals 
appertaining to such species, is a singular fact, and has been strongly insisted on by Dr. 
Hooker, in regard to the Coniferous trees of the southern hemisphere: one would 
naturally have expected, that where circumstances favoured the existence of numerous 
species of a genus, they would likewise have favoured the multiplication of the indi- 
viduals in all or most of such species ; but this, as we here see, has not always been 
the case. 

In the Eocene, Miocene, and Pliocene Tertiary deposits, I know only of two species of 
Scalpellum, and two of Pollicipes, with indications of two or three other species, all distinct 



INTRODUCTION. 7 

from recent forms. It is a rather singular fact, considering the present wide distribution 
of the genus Lepas or Anatifa, and the frequency of the individuals, that not a single 
valve known certainly 1 to belong to this genus, or to any of the closely-allied genera, 
has hitherto been found fossil. 

The oldest known cirripede is, as we have seen, a Pollicipes from the Lower Oolite, and 
it does not differ conspicuously from some of the recent species of the same genus ; so, 
again, the cretaceous Scalp ellum fossula, and the eocene S. quadratum are certainly very 
nearly related to the recent S. rutilum (nov. spec). Loricula alone is a genus perfectly 
distinct from all living Cirripedia ; and I may here add that of the Tertiary Sessile Cirripedes, 
I have hitherto not seen a single new generic form. This persistence of the same genera 
is somewhat remarkable, considering that amongst ordinary Crustacea nearly all the 
Secondary species belong to extinct genera •? it should, however, be borne in mind that 
Limulus has survived from the Palaeozoic period to the present day. The Oolitic, 
Cretaceous, Tertiary, and recent species of Lepadidse are all different from each other. By 
looking at the annexed Table, and putting out of question the species of which the age 
is uncertain, we have five common to two stages of the chalk ; that is assuming for the 
present that the classification of the stages of the chalk commonly used and here followed, 
is correct. Pollicipes glaber is common to three, and, I believe, to four stages. Scalpellum 
arcuatum occurs in the Chalk-marl, and upper Greensand, and therefore this species also 
extends through three stages; but there is a slight difference between the specimens from 
the upper and lower stages, which some authors might perhaps consider specific. If 
fossil cirripedia had, like most recent species, very wide horizontal or geographical ranges, 
then, in accordance with a law now generally admitted, a considerable vertical range in 
some of the species is not improbable. 

I may here observe that I am assured by Professors Forchammer and Steenstrup, that 
the formations of Scania and Westphalia are equivalent to that of Faxoe ; and hence to that 
of Maastricht. I have called these formations the " Maestricht formation" to distinguish 
them from the common upper or white Chalk. 

1 In a mere catalogue, published without descriptions, in the 'Jahrbuch' for 1831, p. 155, by 
Hoenninghaus, Anatifa cancellata is given as a tertiary species : Mr. G. B. Sowerby has stated, in his 
' Genera of Shells,' that he has seen a Tertiary specimen of this genus, but he cannot remember which 
valve it was. 

2 Pictet, Traite Elementaire de Paleontologie, torn, iv, p. 4. 



FOSSIL CIRRIPEDIA. 



TABLE OF THE DISTRIBUTION OF THE SPECIES. 





V 


Faxoe, Scania, 
Maastricht. 


J3 

u 

& 


.a 
O 
u 

o 


a 

M 
13 

J3 


13 
u g 

» a) 


O 


. a 
o » 


So 


3° 


Scalpellum magnum . . . 


* 




















— quadratum . . 


* 




















— fossula .... 





— 


* 
















— maximum . . 





* 


* 
















— lineatum . . . 





— 


— 


* 














— hastatum . . . 





— 


— 





* 












— angustum . . . 


— 


— 


*? 


*? 


*? 












— quadricarinatum . 


— 


— 


— 





* 












— trilineatum . . 





— 


— 





* 












— simplex 


— 


— 


— 








— 





* 






— arcuatum . . . 





— 


— 





* 


— 


* 








— tuberculatum . . 





— 


*? 


#? 


*? 












— solidulum . . . 





* 


















— semiporcatum 


— 


* 


















— (?) cretse .... 





— 


* 
















Pollicipes concinnus . . 


— 
















* 




— ooliticus . . 





— 


— 








— 











* 


— Nilssonii . . . 





* 


















— Hausmanni 
















* 






— politus .... 








— 











*? 








— elongatus . . . 





— 


* 
















— acuminatu8 . . 





— 


— 


* 














— Angelini . . . 





* 


* 
















— reflexus . . . 


* 




















— carinatus . . . 


* 




















— glaber .... 


— 


*? 


* 


* 


* 












— unguis .... 














* 


* 






— validus .... 





* 


















— gracilis .... 


— 


— 


* 


* 














— dorsatus 


— 


* 


















— striatus .... 





— 


* 
















— semilatus . 


— 


— 


*? 


*? 


*? 












— rigidus .... 


— 


— 


— 








— 


* 








— fallax . . 





* 


* 
















— elegans .... 


— 


* 


















— Bronnii .... 












* 










— planulatus . . . 





— 


— 








— 








* 




Loricula pulchella .... 
Total 38 ... . 





— 


— 


* 














4 
* 


9-10 


9-12 


5-8 


5-8 


1 


3-4 


3 


2 


1 



INTRODUCTION. 



NOMENCLATURE OF THE VALVES. 



Figure I. 
CAPITULUM. 



Figure II. 
Scutum of POLLICIPES. 



Apex. 



Occludent 
margin. 




Tergo- 
lateral ! 
margin. 



Basal 
margin. 




Carina. 



Sub-carina. 



Rostral Infra- Carinal 

Latus. median Latus. 

Latus. 

PEDUNCLE. 



N.B. — In the Carina of Scalpellum, 
A is the tectum ; B the parietes ; C the intra-parietes. 





Figure 


III. 




Tergum of pollicipes. 




Apex. 




Occludent 
margin. 


M Wk 
MMssSk 


Upper N 






All 
www 


» \ Carinal 

"J, /margin. 


Scutal 
margin. 


Il» 


Lower 




Basal 
angle. 







Whoever will refer to the published descriptions of recent and fossil Cirripedia, will 
find the utmost confusion in the names given to the several valves ; thus, the valve named 
in the above woodcut, the Scutum, has been designated by various well-known naturalists as 
the " ventral," the " anterior," the " inferior," the " ante-lateral," and the " latero-inferior" 
valve ; the first two of these titles have, moreover, been applied to the rostrum or rostral 
valve of Sessile Cirripedes. The Tergum has been called the " dorsal," the " posterior," the 
" superior," the " central," the " terminal," the " postero-lateral," and the " latero-superior" 
valve. The Carina has received the first two of these identical epithets, viz. the " dorsal" 
and the "posterior;" and likewise has been called the "keel-valve." The confusion, 
however, becomes far worse, when any individual valve is described, for the very same 
margin which is anterior or inferior in the eyes of one author, is the posterior or superior 
in those of another ; it has often happened to me that I have been quite unable even to 
conjecture to which margin or part of a valve an^ author was referring. Moreover, the 
length of these double titles is inconvenient. 

Hence, as I intend to describe all the recent and fossil species, I have thought myself 

b 



10 FOSSIL CIRRIPEDIA. 

justified in giving short names to each of the more important valves, these being common 
to the Pedunculated and Sessile Cirripedes. 

The title of peduncle, which is either naked or squamiferous, requires no explanation ; 
the scales and lower valves are arranged in whorls, which I have called by the botanical 
term of Verticillus. The part supported by the peduncle, and which is generally, though 
not always, in recent species protected by valves, I have designated the Capitulum. 

I have applied the term Scutum to the most important and persistent of the valves, and 
which can almost always be recognised by the hollow giving attachment to the adductor 
scutorum muscle, from the resemblance which the two valves taken together bear to a shield, 
and from their office of protecting the front side of the body. From the protection afforded 
by the two Terga to the dorso-lateral surface of the animal, these valves have been thus 
called. The term Carina is a mere translation of the name already used by some authors, 
of Keel-Valve ; in the genus Scalpellum, in which this valve is taken as typical, 1 have 
found it quite necessary, with fossil specimens, to distinguish the roof (see Woodcut, I,) 
or exterior surface, as the tectum (A) ; the inflected sides, as the parietes (B) ; and in 
several species in the upper half of the valve, the intra-parietes (C) : the expressions of 
apex, basal margin, and inner margin, as applied to the Carina, require no explanation. 
The rostrum has been so called from its relative position to the Carina or keel. There is 
often a sub-carina and a sub-rostrum. 

The remaining valves have been called Latera ; there is always one large upper one 
inserted between the lower halves of the Scuta and Terga, and this I have named the Upper 
Latus or Latera ; the other Latera in Pollicipes are numerous, and require no special 
names ; in Scalpellum, where there are at most only three pair beneath the Upper Latera, 
it is convenient to speak of them {vide Woodcut, I,) as the Carinal, Infra-median, and 
Rostral Latera. 

As each valve, especially amongst the fossil species, requires a distinct description, I 
have found it indispensable to give names to each margin. These have mostly been 
taken from the name of the adjoining valve, (see Woodcut, I.) In Pollicipes the margin 
of the Scutum adjoining the Tergurn and Upper Latus, is not divided (Woodcut, II,) into 
two distinct lines, as in Scalpellum, and is therefore called the tergo-lateral margin ; a 
narrow portion or slip along this side of the valve may be seen (Woodcut, II,) to be formed 
of upturned lines of growth ; this is often of service in classification, and I have called it 
the tergo-lateral slip or segmentum tergo-laterale. In Scalpellum (Woodcut, I,) these two 
margins are separately named Tergal and Lateral. The angle formed by the meeting of 
the basal and lateral or tergo-lateral margins, I call the baso-lateral angle ; that formed by 
the basal and occludent margins, I call, from its closeness to the Rostrum, the rostral 
angle. In Pollicipes the Carinal margin of the Tergum (Woodcut, III,) can be divided into 
an upper and lower Carinal margin. 

That margin in the Scuta and Terga which opens and shuts for the exsertion and 
retraction of the cirri, I have called the Occludent margin. 



INTRODUCTION. 11 

During the periodical growth of the valves, especially when they are thick and massive, 
it happens in several species that the underlying corium deserts their upper ends or umbones, 
which consequently become marked by lines or ridges of growth, as I have called them, 
though perhaps lines of recession would have been more strictly correct. Such valves, 
consequently, have their upper ends projecting from and beyond the capitulum, and are 
said to project freely or lib ere ; this is often more especially the case with the Carina in 
Pollicipes, and in a lesser degree with the Terga. 

From the peculiar curved position which the animal's body occupies within the 
capitulum, I have found it far more convenient (not to mention the confusion of nomen- 
clature already existing) to apply the term Rostral instead of ventral, and Carinal instead of 
dorsal, to almost all the external and internal parts of the animal. Cirripedes have generally 
been figured with their surfaces of attachment downwards, hence I have termed the lower 
margins and angles the Basal, and those pointing in an opposite direction the Upper; 
strictly speaking, the exact centre of the usually broad and flat surface of attachment is 
the anterior end of the animal, and the upper tips of the Terga, the posterior end of that 
part of the animal which is externally visible ; but in some cases, for instance in Coronula, 
where the base is deeply concave, and where the width of the shell far exceeds the depth, 
it seemed almost ridiculous to call this, the anterior extremity ; as likewise does it in 
Balanus to call the united tips of the Terga, lying deeply within the shell, the most pos- 
terior point of the animal as seen externally. 



Class — Crustacea. Sub-class — Cirripedia. 

Family — Lepadidje. 

Cirripedia peduncido jlcxili, musculis instructo : Scutis 1 musculo adductore solummodd 
instructis : valvis cateris, siqaa adsunt, in annulum immobilem haud conjunctis. 

Cirripedia having a peduncle, flexible, and provided with muscles. Scuta 1 furnished 
only with an adductor muscle : other valves, when present, not united into an im- 
movable ring. 

Besides the brief characters here given others might have been added, drawn from the 
softer parts of the animals, but as this Volume treats only of Fossil species, they would 
have been in this place superfluous. Nor have I thought it advisable to give here any 
definition of the Sub-class Cirripedia, or of the Order which contains both the Lepadidae 
and Balanida3, that is the Pedunculated and Sessile Cirripedes ; for the characters would 
likewise have had to be derived almost entirely from the softer parts of the animal. It 
may, however, be worth stating, that by following the metamorphoses of the Cirripedia, it 
can be clearly shown that the capitulum together with the peduncle, in the Pedunculated 
Cirripedes, and that the shell together with the operculum in the Sessile Cirripedes, that 
is the whole of what is externally visible, consists simply of the first three segments of the 
head. In many Crustacea the carapace, formed by the backward production of the three 
anterior rings of the head, covers the dorsal surface of the thorax, and in some it encloses 
the limbs and mouth. This is likewise the case with the Cirripedia, and it is only the 
wonderful elongation of the anterior part of the head, its fixed condition, and the absence 
of external eyes and antennae, which gives to the Cirripedia their peculiar character, and has 
hitherto prevented the homologies of these parts from having been recognised. 2 

1 The meaning of this and all other terms is given in the Introduction at page 9. 

2 Nevertheless, in some Stomapoda, more especially in Leucifer of Vaughan Thompson, the anterior part 
of the head is only a little less elongated, compared with the rest of the body, than in the Cirripedia. 
That accomplished naturalist, M. J. D. Dana (Silliman's 'American Journal,' March, 1846,) has stated 
that " the pedicel of Anatifa corresponds to a pair of antennae in the young :" although the peduncle or 
pedicel is undoubtedly thus terminated, this view cannot, I think, be admitted. In the larva, the part 
anterior to the mouth is as large, in proportion to the rest of the body, as in some mature Cirripedia : this 
anterior part supports only the eyes, antennae, and two small cavities furnished with large nerves, which I 



DSf 



SCALPELLUM. 13 

I may further state, that in the several Orders of Cirripedia such important differences 
of structure are presented, that there is scarcely more than one great character by which all 
Cirripedia may be distinguished from other Crustacea : this character is, that they are 
attached to some foreign object by a tissue or secretion (for at present I hardly know which 
to call it), which debouches, in the first instance, through the prehensile antennae of the 
larva, the antennse being thus embedded and preserved in the centre of the basis. The 
cementing substance is brought to its point of debouchement by a duct, leading from a 
gland, which (and this is perhaps the most remarkable point in the natural history of the 
Class) is part of and continuous with the branching ovaria. When we look at a Cirripede, we, 
in fact, see only a Crustacean, with the first three segments of its head much developed and 
enclosing the rest of the body, and with the anterior end of this metamorphosed head fixed 
by a most peculiar substance, homologically connected with the generative system, to a rock 
or other surface of attachment. 

Genus — Scalpellum. 

Scalpellum. Leach. Journ. de Physique, t. lxxxv, July, 1817. 

Lepas. Linn. Systema Naturae, 1767. 

Pollicipes. Lamarck. Animaux sans Vertebres. 

Polylepas. De Blainville. Diet, des Sc. Nat., 1824. 

Smilium (pars generis). Leach. Zoolog. Journal, Vol. 2, July, 1825. 

Calantica (pars generis). J. E. Gray. Annals of Philosophy, vol. x, (2d series,) 

Aug. 1825. 
Thaliella (pars generis). J. E. Gray. Proc. Zoolog. Soc, 1848. 
Anatifa. Quoy et Gaimard, Voyage de 1' Astrolabe, 1826 — 34. 
Xiphidium (pars generis). Dixon. Geology of Suffolk, 1850. 

Valvis 12 ad 15: Lateribus verticetti inferioris quatuor val sex, lineis incrementi 
plerumque convergentibus ; Subrostrum rarissime adest : Pedunculo squamifero, rarissime 
nudo. 

suspect to be auditory organs ; this part, therefore, I think, must unquestionably consist of the first two or 
three segments of the head : within it, even before the larva moults, the incipient strieeless muscles and 
ovaria of the peduncle can be distinctly traced : immediately after the moult, we see this anterior part 
converted into a perfect peduncle ; and for some time afterwards certain coloured marks, indicating the 
former position of the (so called) olfactory cavities and of the cast-off compound eyes, are still preserved. 
The prehensile antennse are not cast off, for they are fastened down by the cementing substance, and are 
thus preserved in a functionless condition, with their muscles absorbed; after a time even the corium is 
withdrawn from within them. From the above and other coloured marks, and from the antennae being 
preserved, it is easy to point out, in the peduncle of a young though perfect Lepas, the exact point which 
each part occupied in the head of the natatory larva. 

Since the above was written, I find that Loven has taken the same view of the homologies of the 
external parts of the Cirripedia ; in his description of his Alepas squalicola, (Ofversigt of Kongl. Vetens., &c, 
Stockholm, 1844, pp. 192 — 4,) he uses the following words : " Capitis reliquae partes, utin Lepadibus semper, 
in pedunculurn mutatce et involucrum," &c. ; his involucrum is the same as the Capitulum of this work. 



14 FOSSIL CIRRIPEDIA. 

CHARACTERES VALVARUM IN SPECIEBUS FOSSILIBUS. 

Carina angusta, introrsum arcuata, ab apice ad marginem basalem paululum dilatata ; 
parietes valde inflexi, costis manifestis a tecto plerumque disjuncti ; in multis speciebus 
intra-parietibus instrucli : intra-parietes nonnunquam superne producti ultra Umbonem, 
qui fit inde subcentralis : parietum linea incrementi perobliqua. Scuta plerumque 
subconveoca et tenuia, trapezoidea ; marginibus tergalibus later alibusque angulo insigni 
disjunctis. 

Sect. f. Subcarina adest [solummodb species recentes). 

Sect. ft. Subcarina deest. 

A. Valvce quatuordecim : Carina umbone subcentrali. 

B. Valvce duodecim : Carina umbone ad apicem posito. 

Valves 12 to 15 in number. Latera of the lower whorl, four or six, with their 
lines of growth generally directed towards each other. Sub-rostrum 1 very rarely present. 
Pednncle squamiferous, most rarely naked. 

CHARACTERS OF THE VALVES IN FOSSIL SPECIES. 

Carina narrow, bowed inwards, widening but little from the apex to the basal margin, 
having parietes much inflected, and generally separated by distinct ridges from the 
tectum, and having in many species intra-parietes, which are sometimes produced 
upwards beyond the umbo, so as to make it sub-central ; lines of growth on the parietes 
very oblique. Scuta generally only slightly convex and thin, four-sided, the tergal and 
lateral margins distinctly separated by an angle. 

Sect. f. Subcarina present. (This section includes only recent species.) 

Sect. ft. Subcarina absent. 

A. Valves fourteen in number ; Carina with the umbo subcentral. 

B. Valves twelve ; Carina with the umbo at the apex. 

The first of the above two paragraphs contains the true Generic description (here leaving 
out the softer parts), as applicable to recent and, as far as known, to fossil species : the 
second paragraph has been drawn up to aid any one in classifying the characteristic valves, 
when found separated, as is most frequently the case with all fossil Pedunculata. The first or 
proper Generic characters would have been more precise, had it not been for the existence of 
one recent species, the S. villosum (Pollicipes villosus, Leach, Calentica Homii, J. E. Gray,) 
which leads into the next genus Pollicipes. I mention this species in order to confess, that 
had the valves been found separately, and their number unknown, they would certainly 
have been included by me under Pollicipes, although, taking the whole organisation into 
consideration, I have determined to include this species under Scalpellum. I need not 

1 The meaning of this and all other special terms is given in the Introduction at p. 9. 



SCALPELLUM. 



15 



here repeat the remarks made in the Introduction on the great difficulties in classifying the 
recent species, and still more the fossil species of Scalpcllum. I may, however, here state 
that should the 8. vulgare be hereafter kept distinct in a genus to itself, 8. magnum would 
have to go with it. Should a recent species, which in a future work I shall describe under 
the name of S. rutilum, be generically separated, it will probably have to bear the name of 
Xiphidium, from its alliance to the Eocene X. quadratum of Sowerby, to which species the 
cretaceous 8. fossida and several other forms are apparently closely allied. These latter 
species, however, are likewise closely allied to the Scalpellum omatum, which Mr. Gray has 
already raised to the rank of a genus under the name of Thaliella. There are some 
fossil species, as 8. arcuatum, and simplex and solidulum, which I cannot rank particularly 
near any recent forms. Mr. Sowerby founded the genus Xiphidium on the umbo in the 
Carina being situated at the apex, and on its growth being consequently exclusively 
downwards. This is likewise the case with the recent 8. rutilum ; but I shall have 
occasion to show, under 8. magnum, that the upward growth of the Carina in that 
and other species of the genus, depends merely on the intra-parietes, which are present in 
many species, meeting each other and being thus produced upwards. Moreover, in the 
recent 8. omatum, the position of the umbo is variable, according to the age of the 
specimen ; in half-grown individuals being seated at the apex, and in large specimens 
being sub-central, as in 8. vulgare, magnum, and other species. I should have been very 
glad to have retained the genus Xiphidium, but taking into consideration the whole 
organisation of the six recent species, I can only repeat that we must either make six 
genera of them, or leave them altogether, and this latter has appeared to me the most 
advisable course. 

Sexual Peculiarities. — Eor reasons stated in the Introduction, I have kept the genera 
Scalpellum and Pollicipes distinct ; but I may mention, in order to call attention to a point 
of structure which may hereafter be discovered in some fossil species, that I was much influ- 
enced in this decision by some truly extraordinary sexual peculiarities in all six recent species 
of Scalpellum. Scalpellum omatum is bisexual ; the individual forming the ordinary shell, 
is female ; each female has two males (a case of Diandria monogynid), which are lodged in small 
transverse depressions, one on each side, hollowed out, on the inner sides of the Scuta, close 
above the slight depressions for the adductor scutorum muscle ; in 8. rutilum 
(nov. spec.) two males are lodged in the same place on each side, but 
rather in concavities in the valve, than in distinct depressions. As these are ^ 
the two recent species most nearly related to several Cretaceous and Eocene 
forms, we might expect to find similar depressions in some fossil species ; but 
as yet I have not succeeded in distinctly finding such. The male cirripedes 
are very singular bodies ; they are minute, of the same size as the full- 
grown larva; they are sack-formed, with four bead-like rudimental valves at Inside view of tlie Scutum 

° ° in Scalpellum ornatum. 

their upper ends ; they have a conspicuous internal eye ; they are absolutely the ttZttoZ^ for 
destitute of a mouth, or stomach, or anus: the cirri are rudimental and furnished 




16 FOSSIL CIRRIPEDIA. 

with straight spines, serving, apparently, to protect the entrance of the sack : the whole 
animal is attached, like ordinary cirripedes, first by the prehensile antennae, and afterwards 
by the cementing substance ; the whole animal may be said to consist of one great 
sperm-receptacle, charged with spermatazoa ; as soon as these are discharged, the animal 
dies. 

A far more singular fact remains to be told : Scalpellum vulgare is like ordinary cirripedes, 
hermaphrodite, but the male organs are somewhat less developed than is usual ; and, as 
if in compensation, several short-lived males are almost invariably attached on the occlu- 
dent margin of both Scuta, at a spot marked by a fold (not thus caused), as may be seen 
on the inside view of this valve in the fossil S. magnum, which, in all probability, was 
furnished with them. I have called these beings complemental males, to signify that they 
are complemental to an hermaphrodite, and that they do not pair like ordinary males with 
simple females. In Scalpellum vulgare, the complemental male presents only slight specific 
differences from the male of 8. ornatum. It would be foreign to the purpose of this 
volume here to enter on further details ; nor should I have touched on the subject, had I 
not wished specially to call attention to the presence of cavities on the under sides of the 
Scuta above the pits for the adductor muscle. I will only add, that in the other species of 
Scalpellum, the complemental males are more highly organised, and are furnished with a 
mouth and prehensile cirri ; the valves are more or less rudimental in the different species ; 
these complemental males are not always present, and are never attached to young herma- 
phrodites ; when present, they adhere in such a position, that they can discharge their sper- 
matozoa into the sack of the hermaphrodite : their attachment does not affect the form of 
the valves. 1 

Description of Valves. — It will, I think, be most convenient to confine the following 
description to the fossil species of the genus. No one specimen has been found quite 
perfect ; but, judging from analogy, the capitulum was probably formed of fourteen valves 
in S. magnum, and of twelve in the remaining species. These valves are commonly smooth, 

1 Exactly analogous facts are presented, though more conspicuously, by the two species of the genus 
Ibla. Before examining this genus, I had noticed the complemental males on Scalpellum vulgare, but had 
not imagined even that they were Cirripedia. Ibla Cuminyii (as I propose to call a new species collected by 
Mr. Cuming, at the Philippines) is bisexual ; one or two males being parasitic near the bottom of the 
sack of the female. These males are small, are supported on a long peduncle, but are not enclosed in a 
capitulum (such protection being here unnecessary), are furnished with a mouth, ordinary tropin, stomach, 
and anus ; there are only two pair of cirri, and these are distorted, useless and rudimentary ; the whole 
thorax is extremely small ; there is no penis, but a mere orifice beneath the anus for the emission of semen : 
hence Ibla Cumingii is exactly analogous to Scalpellum ornatum. On the other hand, the closely allied 
Australian Ibla Cuvierii, like Scalpellum vulgare, is hermaphrodite, but has, in every specimen*opened by 
me, a complemental male attached to near the bottom of the sack ; this complemental male differs only 
about as much from the male of Ibla Cumingii, as the female J. Cumingii differs from the hermaphrodite form 
of J. Cuvierii. I intend hereafter to give detailed anatomical descriptions and drawings of the males and 
complemental males of Ibla and Scalpellum. 



SCALPELLUM. 17 

but in two or three species are marked with longitudinal ridges ; they are generally rather 
thin ; this, however, is a character which is variable even in the same species. 

Carina narrow, widening but little from the apex downwards, slightly or considerably 
curved inwards, with the umbo seated at the uppermost point : S. magnum, however, 
must be excepted, for in it the umbo is sub-central, and the valve almost angularly bent, as 
will be described in detail under that species. The apex rarely projects freely; but 
this is a variable point in the same species ; the basal margin is either pointed, rounded, or 
rarely truncated. The chief character by which this valve can be recognised, as belonging 
to the genus Scalpellum, is the distinct separation by an angle, (see woodcut, Fig. 1, in the 
Introduction,) often surmounted by a prominent ridge, of the tectum or roof, from the 
parietes, which are either steeply or rectangularly inflected ; the lines of growth on these 
parietes are oblique. A still more conspicuous character is afforded by the part (when 
present), which I have called the intra-parietes ; these give to the valve a pieced appearance, 
and seem let in, to fill up a vacuity between the upper part of the carina and the terga, 
and this is their real office ; they are separated from the true parietes by a ridge, which 
evidently marks the normal outline of the valve. These intra-parietes are flat, and they 
have a striated appearance rather different from the rest of the valve ; and the lines of 
growth on them are extremely oblique, almost parallel to the inner margins of the valve. 

Scuta very slightly convex ; four-sided ; the tergal and lateral margins being divided 
by a slightly projecting point or angle ; and this is the chief character by which the scuta 
of this genus can be distinguished from those of Pollicipes. The umbo is seated at the up- 
permost point, except in S. magnum, and in JS. (?) creta (Tab. I, fig. 1 c, and fig. 11 c), in 
which species the lines of growth, instead of terminating at the angle separating the lateral 
and tergal margins, are produced upwards, so that the valve is added to above the original 
umbo. In S. tuberculatum (fig. 10 c?), the scuta present an intermediate character between 
that in ordinary fossil species, for instance in S.fossula (fig. 4 a), and in 8. magnum and 
creta. The occludent margin is nearly straight, or slightly curved ; both it and the lateral 
margin form nearly rectangles with the basal margin, which is nearly straight. Internally 
the depression for the adductor scutorum is generally, but not always, very plain ; some- 
times the valve is filled up and rendered solid in the upper part above the adductor 
muscle. The apex sometimes projects freely, and is internally marked with lines of growth. 
The internal occludent margin, or edge, is also often marked by lines of growth, and the 
part thus marked, close above the adductor muscle, sometimes becomes suddenly wider ; 
this is caused by some slight change in the position of the animal's body during growth. 

Terga flat, either trigonal or rhomboidal, and, in the former case, sometimes so much 
elongated, with the carinal margin so much hollowed out, as to become almost crescent - 
shaped ; a slight furrow often runs from the upper to the basal angle. Internally, in the 
upper part, there is in some species a little group of small longitudinal ridges, unlike any- 
thing I have seen in recent species, and serving, I apprehend, to give firmer attachment to 
the corium. 



18 FOSSIL CIRRIPEDIA. 

Rostrum unknown in any fossil species ; but judging from recent species, it probably 
existed in all. 

Upper latera known only in three species ; in 8. magnum it is irregularly oval, with 
the umbo central : in 8. quadratum and fossula, five-sided, with the umbo at the upper 
angle : in the eocene 8. quadratum, however, an inner ledge very slightly projects beyond 
the two upper sides, and first indicates a tendency to upward growth. Rostral latera, 
known only in 8. magnum and quadratum, they are transversely elongated, narrow, and 
small. Infra-median latera unknown ; they probably existed only in 8. magnum. Carinal 
latera, known in 8. magnum, quadratum, fossula, solidulum, and maximum; in the first species 
they are transversely elongated ; in the three latter, of an irregular curved shape, and flat. 
In the fossil and recent species, the rostral and carinal latera grow chiefly in a direction 
towards each other ; so that their umbones are close to, or even seated exteriorly to, the 
carinal and rostral ends of the capitulum. Peduncle, calcified scales are known only in 
one species, the 8. quadratum ; but they probably existed in all : the naked peduncle, how- 
ever, of the recent 8. Peronii must make us cautious on this head. 



[A] Valves quatuordecem : Carina umbone suh-centrali. 

1. SCALPELLUM MAGNUM. 1 Tab. I, Fig. 1. 

S. Laterum carinalium et rostralium umbonibus libere (sicut cornud) prominentibus, 
dimidiam seu tertiam partem longitudinis valvarum aquantibus. 

Carinal and rostral latera, with their umbones projecting freely like horns, and 
equalling one half or one third of the entire length of these valves. 

Coralline Crag (lower part). Sutton, Gedgrave, Sudbourne. Mus. S. Wood and Lyell. 

From the close affinity between this species and the recent Scalpellum vulgare, we 
may confidently infer that the capitulum consisted of fourteen valves, which are all pre- 
served in Mr. Wood's collection, with the exception of the infra-median latera and of 
the rostrum. This latter valve would, no doubt, be rudimentary, and it has been over- 
looked by naturalists even in the recent species. The chief difference, excepting size, 
between these two species, is in the form of the rostral and carinal latera, but unfortunately 
these valves are extremely variable. It might even be maintained, with some degree of 
probability, that 8. magnum was only a variety of S. vulgare. The valves of 8. magnum 
are all thicker, stronger, more rugged, and considerably larger than in 8. vulgare. Taking 

1 I have followed Mr. Morris in his Catalogue, in adopting this name from the MS. of Mr. Searles Wood, 
to whose kindness I am greatly indebted for having placed in my hands the whole of his large series of 
valves of this species. 



SCALPELLUM. 19 

the largest scutum, tergum, carina and upper latera in Mr. Wood's collection, they are 
very nearly double the size of the same valves in the largest specimen of 8. vulgare 
seen by me, namely from near Naples, which had a capitulum eight tenths of an inch 
in length ; and they are more than double the size of the same valves in any British 
specimen. Scalpellum magnum probably had a capitulum one inch and a half in length. 

Carina (Tab. I, fig. 1 b and /) abruptly, almost rectangularly bent, with the umbo of 
growth seated just above the bend, at about one third or one fourth of the entire length of 
the valve from the upper point ; form linear, with the lower part slightly wider than the upper. 
Exteriorly the surface is rounded with no central ridge, excepting near the umbo, where the 
narrowness of the whole valve gives it a carinated appearance ; basal margin rounded. Erom 
the umbo two faint ridges run to each corner of the basal margin, separating the steeply- 
inclined parietes from the roof, — a character of some importance in the cretaceous species of 
this genus : outside of these two ridges there are other two ridges, not extending down to 
the basal margin, and separating the parietes from the intra-parietes, which latter being 
united at their upper ends, and produced upwards, form that part of the carina which is 
above the umbo. By comparing the lateral views of the carina of the cretaceous 8. fossula 
(fig. 4 c), and of this species, it will be seen, that the apparently great difference of the 
umbo of growth being either at the apex, or, as in this species, sub-central, simply results 
from the fines of growth of the intra-parietes meeting each other, the valve being thus 
added to at its upper end. The carina of 8. magnum, examined internally, is found often 
to be narrower under the umbo than either above or below it, a character I have not seen 
in the recent 8. vulgare. The lateral width or depth of the valve (measured from the 
umbo to the inner edge) is also greater than in 8. vulgare : this portion is internally filled 
up and solidified. No part of the apex of the valve projected freely. The longest perfect 
specimen which I have seen, is half an inch in length ; but I have noticed fragments 
indicating even a greater size. 

Scuta (fig. 1 c) much elongated, trapezoidal, slightly convex ; umbo placed on the occlu- 
dent margin at about one fourth of the entire length of the valve from the apex, so that the 
valve grows upwards and downwards. Occludent margin straight, slightly hollowed out 
above the umbo, forming rather less than a right angle with the basal margin, which latter 
is at right angles to the lateral margin. The tergal margin is separated from the lateral by 
a slight projection (beneath which the margin is a little hollowed out), and from this pro- 
jection there runs a ridge, often very conspicuous, to the umbo. The part above the ridge, 
stands at rather a lower level than that below it, and the lines of growth on it are generally 
less distinct. This is connected with the fact, as ascertained in 8. vulgare, that the valve, 
during its earliest stage, grows only downwards, the ridge thus indicating the original form 
of the valve and tendency of the lines of growth. On comparing that part of the scuta 
beneath the umbo and ridge, in the present species (Tab. I, fig. 1 c), with the whole valve 
in some other species, for instance in 8. fossula (fig. 4 #), in which the umbo is seated at 
the apex, as it was in the first commencement of growth in 8. vulgare and magnum, it 



20 FOSSIL CIRRIPEDIA. 

will be seen how closely the two valves resemble each other. The scutum of S. tubercu- 
latum (fig. 10 d) is intermediate in its manner of growth between those of 8. magnum and 
fossula. Internally, the impression for the adductor muscle is deep ! on the occludent 
margin, close to the umbo, there is a deep fold, which is connected with the growth of the 
upper part of the valve being subsequent to that of the lower part. There is very little 
difference between this valve and that of S. vulgare ; the upper part, however, appears to 
be always thicker. Length of largest specimen one eighth of an inch. 

Terga (fig. I d) triangular, sometimes approaching to crescent-shaped; flat and thin, 
though the thickness of the valve varies. Carinal margin straight, or very slightly hollowed 
out ; in its upper part there is a barely perceptible prominence marking the limit of the 
upward extension of the carina. Basal angle blunt, rounded ; from it a line, formed by the 
convergence of the zones of growth, runs near and parallel to the carinal margin, up to 
the apex. Occludent margin about equal in length to the scutal ; parallel to the former, 
a slip of the valve is rounded and slightly protuberant, and this portion projects a little on 
the scutal margin. A very small portion, or none, of the apex of the valve projected freely. 
This valve is somewhat narrower, and the scutal margin straighter than in S. vulgare. 

Rostrum unknown, no doubt rudimentary, probably quadrangular. 

Upper latera (fig. 1 e) flat, oval, with the upper half a little pointed ; the lower margin 
shows traces in a varying degree consisting of three sides. The surface, but chiefly of the 
lower half, is faintly marked with stria? radiating from the centre. The umbo lies in the 
middle, and from it two slight ridges, first bending down, diverge on each side. In Scal- 
pellum vulgare this valve (which is very similar in shape to that of 8. magnum) at the first 
commencement of its growth, as with the scuta, is added to only downwards ; and thus the 
two diverging ridges mark the form which the valve originally tended to assume : bearing 
in mind that the basal margin tends to be three sided, if we remove that part of the valve 
above the ridges which have been superadded to the original form, we shall have a five- 
sided valve, essentially like that in the 8. quadratum and S. fossula (fig. 3 e, and fig. 4 d). 

Rostral latera (fig. 1, g to k) elongated, widening gradually from the umbo to the 
opposite end, which is equably rounded : umbonal half free, curling outwards ; the internal 
surface of the other half (A) is nearly flat and regularly oval, with its end towards the umbo 
pointed ; the freely projecting portion varies from nearly one half to one third of the entire 
length of the valve ; but in one distorted specimen it was only one sixth of this length. 
The width, also, of the valve varies (g and h), compared to its length. This valve, com- 
pared with its homologue in 8. vulgare, differs more than any of the preceding valves ; it 
is proportionally larger, and the internal or growing surface is oval, instead of being oblong 
and almost quadrangular ; and the umbonal or freely projecting portion in 8. vulgare is 
only one sixth or one seventh of the entire length of the valve. 
Infra-median latera unknown. 

Carinal latera (fig. 1, I ton) narrow, thick, much elongated, widening gradually from 
the umbo to the opposite end, which is rounded and obliquely truncated. Surface, exteriorly 



SCALPELLUM. 21 

flat ; internally convex. The umbonal, freely projecting portion is sometimes more than 
half, sometimes only about one third, of the entire length of the valve. This portion curls 
outwards and likewise upwards. The degree of curvature and the width (m and n), in pro- 
portion to the length, varies. The upper and lower margins are approximately parallel to 
each other ; the umbonal end of the growing surface is bluntly pointed. This valve differs 
from its homologue in S. vulc/are, in being larger, much narrower in proportion to its length, 
more massive, and with a far larger portion of the umbonal end freely projecting ; also in 
the approximate parallelism of the upper and lower margins, and in the umbonal end of the 
growing surface being pointed instead of square. In 8. vulgare the upper margin is much 
more curled upwards than the lower, and the freely projecting portion is only one fifth of 
the entire length of the valve. 

Taking the largest specimens in Mr. Wood's collection, the freely projecting portions 
of the carinal latera must have stuck out like horns, curling from each other and a 
little upwards, for a length of a quarter of an inch. So again, the much flattened horns of 
the rostral latera, curving from each other, but not upwards, must have projected half an 
inch beyond the probably rudimentary rostrum. The capitulum must have presented a 
singular appearance, represented in the imaginary restored figure (fig. 1 a), with its pair of 
projecting horns at both ends. 

Peduncle ; calcareous scales unknown, but undoubtedly they existed. 

Varieties : the variation in the rostral and carinal latera has already been pointed out. 
In Mr. Wood's collection there are numerous scuta, terga, carinas, and carinal latera, 
from Sutton ; and these are all smaller than those above described, which come from 
Sudbourne, and than some others in Sir C. Lyell's collection from Gedgrave. All these 
places, however, belong (as I am informed by Mr. Wood) to the same stage of the Coralline 
Crag. In the Sutton specimens the carinal latera show the same character as in those from 
Sudbourne, but the carina apparently is not internally so much narrowed in under the 
umbo ; this, however, is a character which is conspicuous only in the larger Sudbourne 
specimens, and anyhow cannot be considered as sufficient to be specific. 

I may take this opportunity of stating, that in Mr. Harris's collection of organic remains 
from the chalk detritus, at Charing, in Kent, I have found the upper part of a carina of 
a very young and minute Scalpellum, which cannot be distinguished from this species; 
but considering the state of the specimen, it would be extremely rash to believe in their 
identity. All the known cretaceous species have the umbo at the apex, so that the Charing 
specimen differs remarkably from its cretaceous congeners. 



22 FOSSIL CIRRIPEDIA. 

[_5] Valvce duodecem : Carina umbone ad apicem posito. 

2. SCALPELLUM QUADRATUM. Tab. I, fig. 3. 

Xiphidium quadratum. Dixon, in Sowerby's Mineral. Conch., Tab. 648 ; Geology of 

Suffolk, Tab. xiv, figs. 3 and 4. 
Pollicipes — ? J. Sowerby. Geolog. Trans., 2d series, vol. v, pi. 8, fig. 5. 

S. tecto parietibusque carince plants, lavibus, simplicibus, margine basalt fere rotun- 
dato ; Lateribus superioribus quinque-lateralibus, lavibus. 

Carina, with its tectum and parietes flat, smooth, and simple ; basal margin almost 
rounded. Upper latera five-sided, smooth. 

Eocene Tertiary. Bognor ; Hampstead. Mus. S. Wood, F. Edwards, N. Wetherell. 

My materials consist of a slab of rock, belonging to Mr. S. Wood, almost made up of 
the valves of this species, of two beautiful specimens in Mr. F. Edwards's collection, and of 
some excellent drawings from Mr. Dixon's specimens by Mr. James de C. Sowerby, in the 
Mineral Conchology. 1 The valves in several of these specimens are nearly in their proper 
positions, though there is not one in which they have not slipped a little. Their relative 
positions are given, I believe nearly correctly, in PI. I, fig. 3 a. Their number I have 
little doubt was twelve. This, however, includes a rostrum, probably almost rudimentary, 
the existence of which I infer only from the analogy of all recent species. Mr. J. Sowerby 
supposed that there were, as in 8. vulgare, four pair of latera (and therefore fourteen valves 
in all), but I conclude, without hesitation, that there were only three pair, as in the recent 
S. rutilum (nov. spec.), to which the 8. quadratum is much more nearly allied than to 
S. vulgare. 

Capitulum : elongated, probably composed of twelve valves. Carina (fig. 3, d, i, k), 
rather narrow, slightly and regularly bowed and widening from the apex to the basal 
margin, which latter is bluntly pointed, or almost rounded ; internally deeply concave ; 
externally with the tectum and parietes flat, and at right angles to each other • — hence the 
carina is square-edged, and its specific name has been given to it. Scuta (fig. 3, b, h) 
oblong, occludent margin slightly arched, forming with the basal rather less than a right 
angle ; tergal margin separated by a just perceptibly projecting point from the lateral 
margin, which latter is very slightly hollowed out; whole valve slightly convex, with a trace 
of a ridge running from the apex to the baso-lateral angle. Internally (A), there is a large 
pit for the adductor scutorum, above which there is a slight depression or fold marked 
with curved lines of growth, and in this depression on each side complemental males 

1 Some small fragments were found by Mr. Wetherell, and are noticed in his Paper in the fifth volume of 
the ' Geolog. Transactions,' entitled "Observations on a Well dug on the south side of Hampstead Heath." 



SCALPELLUM. 23 

were probably attached. Terga (fig. 3 c) triangular, large, flat, basal angle bluntly 
pointed ; apex slightly projecting, as a solid horn ; occludent margin very slightly arched. 
Rostrum unknown; judging from the narrowness of the umbones of the rostral latera, 
it was probably minute or rudimentary. Upper latera (fig. 3 e) large compared with 
the lower valves, flat, five-sided, with the two upper sides the longest ; of the three lower 
sides, that corresponding with the end of the rostral latera is generally (especially in 
young specimens) the shortest. Umbo seated at the uppermost angle ; but in full-sized 
specimens, a narrow ledge has been added, during the thickening and growth of the valve, 
along the two upper margins, and consequently round the apex. Rostral latera (fig. 3/) 
extremely narrow, three or four times as long as wide ; considerably arched, extending 
parallel to the basal margin of the scuta; widening gradually from the umbo to the 
opposite end, which is obliquely truncated in a line (as I believe) corresponding with the 
shortest side of the upper latera ; inner surface smoothly arched ; during growth, the narrow 
rostral half of the valve becomes much thickened, and at the same time added to along its 
upper margin, thus producing a solid, sloping, projecting edge ; umbo slightly projecting. 
Carinal latera (fig. Sg) almost flat, not elongated, of a shape difficult to be described; 
approaching to a triangle, with curved sides, and one angle protuberant. 

Peduncle. The calified scales are apparently large in proportion to the valves of the 
capitulum ; transversely elongated, pointed at both ends, and more or less crescent shaped. 

Affinities. This species was generically separated from Scalpellum by Mr. Dixon, as I 
am informed by Mr. James Sowerby, solely owing to the umbo of growth in the carina 
being at the apex, instead of being sub-central, as in 8. vulgare ; but I need not here 
repeat the reasons already assigned for at present keeping all the recent and fossil species 
under the same genus. In the umbo of growth, in the carina and scuta being seated at 
their upper ends, in the square form of the carina, in there being only three pair of latera, 
and in the large size of the upper latera, this eocene species is much more closely allied to 
8. rutilum (nov. spec., of which the habitat is unfortunately not known,) than to any other 
recent species. In some respects, however, I may remark, 8. rutilum is even more closely 
related to certain cretaceous forms. To 8. ornatum, the nearest recent congener of 8. rutilum, 
the present species is allied by the narrowness of the rostral latera, and by the large size 
and peculiar shape of the scales on the peduncle : the carinal latera perhaps rather more 
resemble those of 8. vulgare than of any other recent species. Certainly, all the affinities 
in 8. quadratum point to 8. rutilum, ornatum, and vulgare, and these three recent species 
are characterised by having males or complemental males attached to the sides of the orifice 
of the sack, whereas, in the other species, they are elsewhere attached ; hence it is that I 
believe that males were probably lodged in the slight depressions described on the inner 
sides of the scuta ; but the depression is not here nearly so distinctly developed as it is 
in the recent 8. ornatum, and more resembles the fold on the occludent edge of the valve 
in 8. vulgare : I must add that folds of this nature do not necessarily imply the presence 
of males. 



24 FOSSIL CIRRIPEDIA. 

3. SCALPELLTJM FOSSULA. Tab. I, fig. 4. 

Pollicipes maxtmus. J. Sowerby. Min. Conch. Tab. 606 (a tergum), fig. 3. 

S. carina intra-parietibus instructd ; tecto utrinque costis magnis, tumidis, superne 
planatis, marginato ; margine basalt obtuse acuminato. Lateribus superioribus quinque- 
lateralibus ; costis duabus modicis ab apice ad marginem basalem continuatis. 

Carina, having intra-parietes, with the tectum bordered on each side by large, pro- 
tuberant, flat-topped ridges ; basal margin bluntly pointed ; upper latera five sided, with 
two slight ridges extending from the apex to the basal margin. 

Upper Chalk. Norwich ; Northfleet, Kent. Mus. Fitch, J. de C. Sowerby, Wetherell. 

General Remarks. My materials consist of two specimens, belonging to Mr. Fitch, 
most kindly lent me for examination ; in which, taken together, the scuta, terga, carina, 
upper and carinal latera, are seen almost in their proper places. In Mr. J. Sowerby's 
collection there is a single scutum, also, from Norwich. From analogy with the eocene 
8. quadratum and the recent S. rutilum, I have little doubt that there were only three pair 
of latera ; and that, probably, there was a rostrum. With respect to the exact position of 
the carinal latera, there is, as also in the case of the 8. quadratum, some little doubt. 

Capitulum narrow, elongated, probably composed of 12 valves, which are moderately 
strong, and apparently closely locked together. The length of the capitulum in the largest 
specimen was 1*1 of an inch. 

Carina (fig. 4, c, g, h) strong, moderately bowed, extending far up between the terga, 
almost to their upper ends ■ rather narrow throughout, gradually widening from the apex 
to the base ; lines of growth plain ; no portion projects freely. The tectum or central 
portion is slightly arched, subcarinated, and bounded on each side by flat-topped, pro- 
tuberant ridges : the tectum terminates downwards in a blunt point (the two margins 
forming an angle of rather above 90°), which projects beyond the bounding ridges ; the 
tectum and the two bounding ridges all widen gradually from the apex towards the 
base. The parietes are channelled or concave ; they do not extend so far down as 
the ridges bounding the tectum. In the upper half of the carina, we here first see 
the additional parietes, or intra-parietes, which appear as if formed subsequently to 
the other parts, and let in between the ordinary parietes of the carina, and the terga. 
It has been already shown, under 8. magnum, that it is the intra-parietes produced 
upwards, which causes in that and some other species the umbo of the valve to be sub- 
central. 

Scuta (fig. 4, a,f) oblong, the basal margin only slightly exceeding half the entire length 
of the valve ; valve strong, rather plainly marked with lines of growth ; basal margin at 
nearly right angles to the occludent margin ; tergal margin separated by a slightly-projecting 



SCALPELLUM. 25 

point from the lateral margin, which in the lower half is slightly protuberant ; tergal 

margin straight, with the edge thickened and slightly reflexed. A distinct, square-edged 

ridge (therefore formed by two angles) runs from the umbo to the baso-lateral angle, which 

is itself obliquely truncated. Internally (/), there is a large and deep pit for the adductor 

scutorum. Terga (fig. 4 b) triangular, flat, large, fully one third longer than the scuta ; 

basal half much produced ; basal angle pointed ; from it to the apex or umbo there runs 

a narrow, almost straight furrow, at which the lines of growth converge — it runs at about 

one third of the entire width of the tergum (in its broadest part) from the carinal margin. 

Parallel to the occludent margin, and at a little distance from it, there runs a wide, very 

shallow depression up to the apex. The scutal margin is not quite straight, about a third 

part, above a slight bend corresponding with the apex of the upper latera, being slightly 

hollowed : from the above bend a very faint ridge runs to the apex of the valve. Upper latera 

(fig. 4 d) large, flat, with five sides, of which the two upper are much the longest ; the 

basal side is next in length, and the scutal side much the shortest. As far as I can judge 

of the positions of the lower valves, with respect to the upper latus, I believe, that the 

rostral latera, probably, abutted against the shortest of the three lower sides ; that the 

carina ran along the one next in length, and the carinal latera along the middle basal 

side, which I suppose extended in an oblique line, and not parallel to the base of the 

capitulum : the two upper long sides no doubt touched the scuta and terga. The umbo 

of growth is at the apex ; there is, however, a trace of a projecting ledge added round the 

upper margins during the thickening of this upper part of the valve. Two slight ridges run 

from the apex to the two corners of the middle of the three lower sides. Carinal latera 

(fig. 4 e) : these are not quite perfectly seen : the umbo forms a sharp point, whence the 

valve rapidly expands and curves apparently downwards and towards the upper latera. 

Near one margin there is a very narrow furrow, and on the other a wide depression, both 

running and widening from the umbo to the opposite end, which is slightly sinuous. I 

imagine these carinal latera occupied a nearly triangular space between the middle of the 

three lower sides of the upper latera and the basal portion of the carina. Rostral latera, 

rostrum and peduncle unknown ; the rostral latera must have been very narrow. 

Affinities. — In the shape and manner of growth of the scuta, and more especially of 
the upper latera, this species is certainly more closely allied to the eocene 8. quadratum, 
than to any other species ; but in the peculiar characters of the carina, it is nearer to the 
recent 8. rutilum ; we have previously seen that the nearest congener to 8. quadratum is 
this same S. rutilum. The most conspicuous diagnostic character of this species is derived 
from the peculiar form of the carina, — its tectum being bounded by a rounded ridge on 
each side. The square-edged ridge running from the apex to the baso-lateral angle of 
the scuta is a trifling, but I believe, a diagnostic character. If I am right in placing 
S. rutilum in the genus Scalpellum, and I think there can be no doubt of this, considering 
the characters of its complemental male, then there can be no question that the present 
species belongs to the same genus. 

d 



26 FOSSIL CIRRIPEDIA. 



4. SCALPELLUM MAXIMUM, Tab. II. fig. 1 10. 

Pollicipes maximtjs. J. Sowerby. Min. Conch., tab. 606, solummodo, fig. 4 et fig. 6. 
N. B. — Fig. 3 est Tergum S. fossulce, et fig. 5 alia species 
ignota. 

— maximus. Steenstrup. Kroyer Tidsskrift, b. ii, pi. v, figs. 17, 18. 

— medius. Steenstrup. Kroyer Tidsskrift, b. ii, pi. v, figs. 13, 13*, 33. 

— sulcatus. J. Sowerby. Min. Conch., pi. 606, fig. 2, sed non fig. 1 et 7. 

5. carina intra-parietibus instructd ; tecto subangulari vel subcarinato ; margine basalt 
rectangulariter acuto ; totd valvd plus minusve introrsum arcuatd, sed margine inferno fere- 
recto ; tecto 1 transverse plus minusve convexo j superficie pane lavi, striis paucis obsoletis 
longitudinalibus elevatis ; tectum, parietes, et intra-parietes inter se separantur costis plus 
minusve prominentibus. 

Carina having intra-parietes, with the tectum slightly angular or subcarinated, basal 
margin rectangularly pointed : whole valve more or less bowed inwards, but with the inner 
margin nearly straight ; tectum, in a transverse line, more or less convex ; surface nearly 
smooth, with a few faint longitudinal raised striae ; more or less prominent ridges separate 
the tectum, parietes, and intra-parietes from each other. 

Upper Chalk, Norwich (common), Mus. Fitch. Northfleet (single spec), Kent, Mus. J. Sowerby. 
Upper Chalk, Charing, Kent, Mus. Harris. Scania, and Quedlingburg in Westphalia, Mus. University, 
Copenhagen. Cyply bei Mons, Belgium, Mus. Brit. Gehrden Hanover, oberer Kreidemergel, Mus. 
Dunker and Roemer. 

I have had far more difficulty in making up my mind regarding this the commonest 
cretaceous species, than with all the other fossil pedunculated cirripedes. From reasons pre- 
viously stated, I have in this genus, when only separate valves have been found, taken the 
carina as typical. Comparing ordinary specimens of the carina of Scalpellum maximum and 
var. sulcatum, such as those figured in the ' Mineral Conchology,' I should certainly have 
considered them quite distinct, had not an examination of Mr. Fitch's fine collection 
from Norwich, together with several other specimens, shown me that there are intermediate 
forms which it is scarcely possible to class. Again, had I not seen a particular carina of 
S. maximum var. cylindraceum, in which the upper part displays a different character from 
the lower in the same individual valve, I should have unhesitatingly received it as a 
species, instead of, as I now do with certainty, as a mere variety. I feel, moreover, very 
great doubts whether the S. lineatum be a species, or merely another variety of 8. maximum ; 

1 For an explanation of this and all other terms, see the remarks on nomenclature and woodcuts in the 
Introduction, page 9 and 10. 



SCALPELLUM. 27 

its distinctive characters are extremely slight ; but they do not blend away by any inter- 
mediate forms hitherto seen by me. Looking only thus far, it would have been natural to 
have classed, without any doubt, all the carinae as varieties of 8. maximum, but in the 
same Norwich beds, from which Mr. Fitch obtained his fine series of carinas, there are 
scuta and terga, which undoubtedly belonged to the genus Scalpellum, and which, from 
being of equally large size, nearly equally numerous, and having a similar state of surface 
with the above carinae, I believe belonged to them : but both the terga and scuta 
present a more remarkable range of variation than do even the carinas. In the case 
of the terga, at one extreme of the series, I did not even at first recognise the valve to 
be a tergum ! yet the forms so blend together with very short intervals, that I cannot 
specifically separate them. Terga of the two extreme forms come, also, from the same 
localities in Scania. In the case of the scuta there are three distinct forms in Mr. Pitch's 
collection, which I should certainly have considered as specifically distinct, had I not 
been led from studying the carinas and terga to believe that this species varies much : 
moreover, the chief point of variation in the scuta, namely, in the character of the under 
surface of the upper part, I conceive to be, in some degree, in connection with one chief 
peculiarity in the terga, namely, the varying prominence of their occludent margins. 
Although I have not seen any other instance of so much variation in the scuta; yet 
I believe that I have taken the most prudent and correct course in describing them as 
mere varieties. From the more frequent coincidence of the carina, described as that of the 
true maximum, with the Varieties I of the scuta and terga, I believe that these valves 
belonged to the same individuals : with respect to the two other varieties, I have hardly 
any grounds for conjecturing which belonged to which. It is most unfortunate that 
not a single specimen of this species seems, hitherto, to have been found with all, or 
even a few, of its valves embedded together. 

In giving names to the varieties, as judged of by the Carinae, there is a difficulty in 
nomenclature ; for the carina of 8. maximum and of 8. maximum, var. sulcatum, are appa- 
rently almost equally numerous in the Norwich beds ; and might either be taken as 
typical of the species ; I have chosen the former name, simply as having been more com- 
monly used, and from this form having been apparently most widely distributed. I have 
described under it the original carina of Pollicipes maximus of J. Sowerby, and all the 
other valves, which I have reason to suppose belonged to this species. The other carinae, 
however, as being in this genus the typical valve, are described under separate subordinate 
headings ; the description of 8. maximum, var. sulcatum, being given from Mr. Sowerby's 
original specimen. Under the typical 8. maximum, I indicate as far as able, to which 
carinae the varieties of the scuta and terga, there described, probably belonged. 



28 FOSSIL CIRRIPEDIA. 



Scalpellum maximum, var. typicwm. Tab. II, figs. 1, 4, 5, 8. 

S. carina introrsum leviter arcuatd, latitudine valvce altitudinem super ante ; tecto 
transverse leniter arcuato ; parietibus intra-parietibusque angustis, superficie pane 
Icevi. 

Carina slightly bowed inwards ; width of valve greater than the depth ; tectum flatly 
arched transversely ; parietes and intra-parietes narrow ; surface nearly smooth. 

Carina, Tab. II, fig. 1. In this, the typical variety, the carina is very slightly bowed 
inwards, widening gradually downwards from the apex, of which no part projected freely \ 
walls rather thin ; tectum very flatly arched, not sub-carinated ; basal margin rectangu- 
larly or rather more acutely pointed ; parietes very slightly concave, splaying outwards, 
narrower than one side of the tectum, separated from it and from the intra-parietes by 
rounded ridges ; intra-parietes narrow, not extending baseward so far as the basal margin 
of the parietes ; width of valve measured from marginal edge to edge, considerably greater 
than the depth, measured in the same place from the central crest to either marginal edge ; 
but the width compared with the depth varies a little : inner margin of valve nearly 
straight. Length of longest specimen (Mm. Fitch) nearly 1^ inch. This variety in the 
Norwich beds seems about equally common with var. sulcatum, but the former alone is 
found in Hanover and in Scania, excepting that in the latter country some specimens indi- 
cate a passage to the var. cylindraceum. 

Scutum, Tab. II, fig. 8. In Mr. Fitch's collection there are three left-hand valves of a 
Scalpellum, which, from their size and smoothness, I have no doubt belonged to this species, 
and from then thinness, probably to the variety of carina considered as typical under the 
simple name of S. maximum ■ valve unusually thin and little convex ; trapezoidal, with 
the apex less produced than is usual in the genus ; broad in proportion to its length, the 
basal margin being '66, and the occludent margin '98 in length, therefore the breadth 
equals two thirds of the length. Basal margin (just perceptibly hollowed out) forming less 
than a right angle with the (just perceptibly outwardly arched) occludent margin, and 
forming an almost exact rectangle Avith the lateral margin ; the latter meets the tergal 
margin at an angle of about 135°. The edge of the tergal margin is thickened and 
slightly reflexed ; the upper part of the lateral margin is in some specimens a little bowed 
inwards. The baso-lateral angle is rounded and just perceptibly protuberant ; no ridge 
or furrow runs from it to the apex. Internally the depression for the adductor muscle is 
singularly shallow (fig. 8 c); a very small portion of the upper part of the valve projected 
freely ; the internal surface of the valve, above the pit for the adductor muscle, has not 
been thickened, and is therefore slightly concave or almost flat. The internal occludent 
edge in the upper part is only a very little widened, and is flat ; on the tergal margin, 



SCALPELLUM. 29 

a narrow ledge of about equal width with the occludent edge, marked likewise with lines 
of growth, must have overlapped the tergum. Largest specimen 1*15 in length. 

Scutum, Var. II, Tab. II, fig. 9. This valve is narrow, moderately convex, with the 
upper portion much acuminated; the tergal margin is somewhat hollowed out, and is 
bordered by a narrow smooth slip, (as in the scutum of 8. arcuatwn,) which is simply 
formed by the thickening from within of the upper part of the valve ; this slip does not 
reach to the uppermost point. The occludent margin is somewhat arched, at nearly right 
angles to the basal margin ; lateral margin forming an angle a very little above a right 
angle with the basal margin. A conspicuous, curved, angular ridge runs from the 
apex to the baso-lateral angle, (which is not at all protuberant,) and divides the valve 
obliquely into two almost equal halves. Surface just perceptibly striated, finely and 
longitudinally. Internally there is a deep pit for the adductor scutorum, which is situate 
low down in the valve ; the inner occludent edge in the upper part of the valve {b), above 
the adductor scutorum, widens suddenly, and is formed into a furrow, which, however, 
I do not believe to have had any functional importance ; the central internal surface 
of the valve, above the pit for the adductor muscle, is somewhat prominent; and a 
quite small, almost flat, portion of the tergal side is marked by lines of growth, showing 
where it overlapped the tergum. Altogether there is a considerable resemblance between 
this valve, both externally, and more especially internally, and that of the Pollicipes 
Angelini. Prom the valve being acuminated, with the upper part rather solid, and from 
the surface being just perceptibly striated, it more probably belonged to var. sulcatum than 
to the typical 8. maximum. 

Scutum, Var. Ill, Tab. II, fig. 10. This third variety, of which the specimen is a fine 
large one, is about intermediate in outline or •acumination between the first and second 
varieties : the tergal margin is thickened and reflexed as in the first, and is not bordered 
by a smooth narrow slip as in the second variety. There is no distinct angular ridge, as 
in the second variety, running fromi the apex to the baso-lateral angle. Internally the 
differences are more conspicuous ; the depression for the adductor muscle is pretty well 
developed ; a large portion of the upper part of the valve projected freely ; the internal 
occludent edge, above the adductor-depression, becomes greatly widened and deeply 
hollowed out, but yet the furrow I believe, as in Var. II, to be of little or no functional 
importance, and merely a consequence of the internal thickening of the central upper part 
of the valve ; on the tergal side a wide ledge shows the extent to which that margin 
overlapped the tergum. The internal surface of the valve, above the adductor-depression, 
is filled up solid and is exceedingly prominent, as is the ridge extending from it to the 
apex ; this ridge, from the unusual width of the internal occludent edge, is pushed over to 
the tergal side of the valve. 

Professor Steenstrup has sent me two small scuta, collected by M. Angelin at Kopinge 
and Balsberg, in Scania, which come near to the Third variety; the internal furrow, however, 
along the occludent margin, is much narrower, deeper, and oblique, so that it is partly 



30 FOSSIL CIRRIPEDIA. 

covered by a lateral projection of the central portion : a tolerably distinct ridge runs from 
the apex to the baso-lateral angle. Amongst the several specimens from Hanover sent me 
by Drs. Dunker and Roemer, the scuta all belong to the First variety. 

I believe all these differences in the scuta of the three varieties ensue partly from the 
varying acumination of the upper part, and consequently of the extent to which the apex 
projected freely, but chiefly from the degree to which the upper part of the valve above 
the adductor muscle has been internally thickened. In the first variety the upper part is 
simply concave, and the pit for the adductor very shallow ; in the third variety, the same 
upper part is highly prominent, and apparently as a consequence the internal occludent 
edge is deeply furrowed ; the pit for the adductor muscle is deepest in the second variety. 

The above differences would perhaps affect the outline of the terga, but I am not able to 
follow the precise manner ; nor should I have thought them sufficient to have produced the 
amount of variation presently to be described in the terga ; but possibly other scuta may vary 
still more. At first I concluded that the upper part of the inner occludent edge, which in 
Var. Ill is deeply furrowed, received in it the occludent edge of the tergum (as the furrow 
on the tergal side of the apex of the scutum receives the edge of the tergum in the recent 
Pollicipes mitetta), but this on consideration I do not think can possibly be the case, 
although it would amply account for the variation in the terga. 

Terga. I have seen great numbers of these valves ; eight specimens are in Mr. Fitch's 
collection from Norwich ; one is figured by Mr. J. Sowerby in the ■ Min. Conch.,' (PI. 606, 
fig. 6,) and they are numerous in the collection from Scania and Hanover. These valves, 
which, as stated in the preliminary remarks, present a most remarkable amount of variation, 
will be best described under three distinct heads. 

Variety I. Tab. II, fig. 5. This valve, from its greater width and smoothness, 
compared with the other varieties, perhaps belongs to the typical S. maximum. Surface 
smooth, with a mere trace of some longitudinal strise, sub-rhomboidal, elongated, with the 
apex much produced and curled towards the carina ; nearly flat ; the occludent margin 
arched, nearly equal in length to the scutal margin ; upper carinal margin hollowed out, 
about half the length of the lower carinal margin ; the occludent and upper carinal margins 
meet each other at a very small angle, making the apex almost horn-like ; from it to the 
bluntly pointed basal angle, a slight rounded ridge, and on the carinal side of it a slight 
furrow, (both becoming less plain towards the lower part of the valve,) extends. As seen 
internally, the thickness of the valve, in its upper part, varies ; a rather large upper part 
projects freely. A rim along the occludent margin is rounded and slightly protuberant, 
with a slight depression in the valve parallel to it. Length of the largest specimen 1*2 of 
an inch. 

This variety is found commonly near Norwich, in Scania, and Hanover. 

Tergum. Variety II. Tab. II, fig. 6. The valve in this variety (from near 
Norwich) is much elongated, sub-triangular, approaching to crescent-shaped; lines of 
growth conspicuous, with a few very faint longitudinal striae. Carinal margin not (or 



SCALPELLUM. 31 

barely) distinguishable into an upper and lower portion ; the whole being nearly straight, 
or very slightly concave. Apex extremely produced, narrow, and horn-like; curled 
towards the carina; apparently (for the apex is broken) a considerable portion was 
thickened, and must have projected freely. Occludent margin slightly arched, about equal 
in length to the scutal margin, which latter in the lowest part is curved and projects a 
little. Basal angle bluntly pointed. A rounded ridge (with a mere trace of a furrow on 
its carinal side), almost disappearing in the lower part of the valve, runs from the apex 
to the basal angle in a slightly curved course, strictly parallel to the carinal margin. The 
rim of the valve along the occludent margin is rounded and strongly protuberant, and, 
parallel to it, the surface is considerably depressed. Length of valve, when perfect, 
12 of an inch. This variety differs from the first in the much greater straightness of the 
carinal margin, in the occludent rim being more protuberant, and in the scutal margin 
not being quite straight. One specimen presented a decidedly intermediate form, though 
rather nearer to the first than to the second variety. 

Tergwm. Var. III. Tab. II, fig. 7. The valves of this variety, of which I have seen 
five specimens, were for a long time quite unintelligible to me, and I at first even thought 
that perhaps they were rostral latera, but I now find that in outline, though not in 
general appearance, owing to their great thickness, they closely resemble the terga of S. 
magnum. One of the four specimens is almost exactly intermediate between the variety 
last named and that now to be described ; hence there can be no doubt that they are really 
terga. The chief characteristic of the valves of this variety is their narrowness, and the 
solidity of their upper ends, which, together with a point of structure presently to be men- 
tioned, makes me think it likely that they belonged to the individuals which possessed a 
carina, hereafter to be described under the name of S. maximum, var. cylindraceum. Valve 
smooth but with the lines of growth plain, extremely narrow, almost crescent-shaped ; 
carinal margin considerably more concave than in Var. II, with a barely perceptible promi- 
nence in the upper part, marking the commencement of the freely projecting portion, 
and probably the point of upward extension of the carina. The occludent margin is 
arched, and is equal in length to the straight scutal margin. From the apex there runs a 
fine furrow (instead of a ridge and furrow, as in Vars. I and II,) to the basal angle, nearly 
parallel to the carinal margin, but almost blending with it in the lower part of the valve. 
The upper freely projecting portion is much thickened, and rendered almost horn -like, but 
to a variable extent ; owing to this the width of the valve in the upper part also varies. 
In the specimens most characteristic of the present variety, the rim of the valve along the 
occludent margin is not at all, or barely, protuberant, nor is there any plain depression 
parallel to the occludent margin : in the intermediate specimen, however, above alluded to, 
the rim is protuberant and there is a plain depression, though both much less conspicuous 
than in the tergum of Var. II. On the internal surface of the upper freely projecting 
part, (marked with lines of growth,) there can be observed in two specimens a slight 
and variable longitudinal depression ; judging from what occurs in the recent genus 



32 FOSSIL CIRRIPEDIA. 

Lithotrya, and from what may be faintly seen in the tertiary Pollicipes carinatus, and even 
in some specimens of P. mitetta, I believe that this structure indicates that the upper freely 
projecting portion of the carina had its inside filled up and rendered prominent, which we 
shall see is the case with the carina of the variety cylindraceum. Length of largest 
specimen, eight tenths of an inch. 

This variety is found at Norwich, in Scania, and at Cyply bei Mons, in Belgium. 

Amongst the Scanian specimens from Kopinge (where the carina of the true iS. maxi- 
mum is commonly found) there are some terga differing from the variety just described, 
only in having the lower part of the valve less produced ; and more especially in having 
on the internal surface of the upper part a smooth prominent ridge, lying rather nearer to 
the occludent than to the carinal margin, and therefore in exactly the same position in 
which a little group of small, sharp, longitudinal ridges occurs in the terga of S. arcuatum 
and of some other species. I am surprised at such a point being variable, but I cannot 
doubt that this valve belongs to the same species. I may add that it was this trifling 
point of structure, which first led me to suspect that these singular crescent-shaped 
valves were really terga. Finally, I may remark, that when all the ten terga now described 
are placed in a row, it is scarcely possible to doubt that they form merely varieties of the 
same species. 

Carinal latus, Tab. II, fig. 4. This is the only valve which remains to be described, 
for neither the rostrum nor rostral latera are as yet known. It was found at Kopinge, in 
Scania, where the carina of the true S. maximum occurs abundantly ; it was sent to me by 
Professor Steenstrup, who attributed it to this species. I have also seen a specimen from 
Hanover, where the carina of the true S. maximum is also found, and another small 
specimen from Charing, in Kent. Valve thin, of an irregular shape, sub-triangular ; flat, 
except at the umbo, which projects outwards, owing to a ledge formed beneath and round 
it ; carinal margin very slightly convex, with a linear furrow parallel to it, between which and 
the edge the lines of growth are abruptly upturned ; lower margin considerably convex « 
upper margin slightly concave, with a slight depression parallel to it, between which and 
the edge the lines of growth are rectangularly reflexed towards the umbo. The two Scanian 
specimens differed slightly in outline ; chiefly with respect to the projection of the ledge 
round the umbo. Width of largest specimen one quarter of an inch. This valve unmis- 
takeably resembles the homologous valves in S. quadratum and fossula, but can be 
distinguished from both ; the end opposite the umbo is much less produced than in S. 
quadratum ; the whole valve is wider, and the furrows much less developed, than in 
S. fossula, to which it comes nearest. 

Affinities. Before describing the several varieties as characterised by their carina?, I will 
offer a few remarks on the affinities of this, the most common and widely distributed species 
of all the cretaceous pedunculated cirripedes. Mr. James Sowerby at first naturally 
described it as a Pollicipes ; quite lately in Mr. Dixon's work he has considered it as 
belonging to the same genus with his eocene Xip/iidium quadratum and our Scalpellum 



SCALPELLUM. 33 

quadratum. Still closer is the affinity with the cretaceous 8. fossula ; the carinas of both 
have intra-parietes; the tectum is distinct from the parietes, which latter are either 
channelled or concave ; the trapeziform scuta of 8. quadratum, fossula, and maximum, are 
unmistakeably alike, and even more striking is the resemblance of the carinal latera ; 
there can be no doubt of these three species belonging to the same genus, and having 
the same number of valves, namely, as I have shown under 8. quadratum and fossula, 
probably twelve. 

Geological History. This species, with its varieties cylindraceum and sulcatum, is very 
common in the Upper Chalk strata of Norwich ; I have seen one specimen from the Upper 
Chalk of Northfleet, in Kent. It is common in the sandstone beds of Scania, which I am 
assured by Professor Forchhammer, are without doubt equivalent with the Faxoe beds, and 
therefore belonging to a stage above our flinty chalk. I have seen, also, one specimen, 
belonging, I believe, to this species, from the same stage in Westphalia ; and another from 
Belgium ; it is also common at Gehrden, in Hanover, in the ' Oberer Kreidemergel' of 
Roemer. 

SCALPELLUM MAXIMUM, VAR. CYLINDRACEUM. Tab. II, fig. 2. 

S. parte superiore carina libere prominente, parte interiore intra-parietibus rotundatis, 
inflexis, itd repletd, ut pane cylindrica flat ; superficie externa lavi, tecto parietibusque pane 
confluentibus. 

Carina, with the upper portion projecting freely, and with the inside filled up by the 
rounded inflected intra-parietes, so as to be almost cylindrical; exterior surface smooth, 
with the tectum and parietes almost confluent. 

Amongst the specimens from Norwich, two differed from the others in being a little 
more elongated and smoother, in the parietes becoming almost confluent, low down on the 
valve, with the tectum, and in the intra-parietes being very little developed. On tne 
internal face this variety presents its most remarkable character ; for a large upper portion 
of the valve must have projected freely, and the intra-parietes, instead of forming a 
thin wall on each side, are thickened, rounded, and turned inwards, so as almost to meet, 
and thus to fill up the original concavity of the valve. Hence a section (fig. 2, c) of 
the upper part, some way below the apex, is almost cylindrical, or more strictly oval 
with the longer axis in the longitudinal plane of the animal, with either a wedge-formed 
hollow, or a mere, almost closed, cleft on the under side, penetrating not quite to the 
centre of the solid valve. The two specimens differ, one in being in a transverse line 
exteriorly much depressed, the other highly arched or convex, and internally still more 
conspicuously in the degree to which the intra-parietes have filled up the upper part. In 
one of the specimens there is even a difference on the opposite sides of the same individual 
valve. Notwithstanding these varieties, I should have much hesitated to have ranked 

e 



34 FOSSIL CIRRIPEDIA. 

these peculiar carinse under 8. maximum, had not the upper part in one specimen actually 
retained all the usual characters of 8. maximum, the precise line where the manner of 
growth had changed, being distinctly visible. It is represented in Plate II, fig. 2, a and b. 
Amongst the Scanian specimens, some make an approach to this variety. 



SCALPELLUM MAXIMUM, VAR. SULCATUM. Tab. II, fig. 3. 

Pollicipes sulcatus. J. Sowerby. Min. Conch., pi. 606, solummodo, fig. 2. Fig. 7 
fortasse Carina P. Angelini. Fig. 1, Tergum fortasse P. 
striati. 1 

8. carina introrsum valde arcuatd, sub-carinatd ; valva latitudine circa dimidium 
altitudinis cequa?ite, tecto transverse prarupte arcuate-; parietibus intra-parietibusque 
latiusculis. Apice solide repleto, libere paululum prominente ; superjicie externa striis 
paucis, rotundatis, ad alterum' vel utrumque latus costarum duarum tectum et parietes 
separantium. 

Carina considerably bowed inwards, subcarinated ; width of valve about half of the 
depth ; tectum in a tranverse line, steeply arched ; parietes and intra-parietes rather 
wide; apex filled up solid, and projecting freely a little; exterior surface with a few 
rounded striae on either one or both sides of the two ridges which separate the tectum and 
parietes. 

Having had the advantage of seeing Mr. J. Sowerby 's original specimen, the valve 
now to be described is certainly that figured by him as Pollicipes sulcatus. As already 
stated, certain specimens of this variety differ strikingly from the carinae typical of 8. 
maximum ; whereas others, from the same formation and locality, are so intermediate 
that they can, with difficulty, be arranged on either side : this is also the case with 
one from Cyply bei Mons, in Belgium. This variety is common in the Upper Chalk 
of Norwich. 

In a well-marked specimen of this variety, the chief distinctive characters, as contrasted 
with the true 8. maximum, consist in the tectum being more steeply arched, in the depth 
of the valve being much greater than the width, in the intra-parietes and parietes being 
more developed, in the whole valve being more bowed inwards, in the walls being 
thicker and apex filled up solid, in the surface having a few fine raised lines on each 
side of the ridge separating the tectum and parietes, and, lastly, in the tectum being 
sub-carinated. 

1 If I am correct in considering the carina of P. sulcatus to be only a variety of that of S. maximum, 
the tergum figured by Mr. Sowerby as belonging to his P. sulcatus cannot so belong ; for it does not at 
all resemble the homologous valve of S. maximum. I believe from the character of the ridge running from 
the apex to the basal angle, that it belonged to a Pollicipes, which must have been coarsely striated longi- 
tudinally, and therefore I have provisionally described it under Pollicipes striatus. 



SCALPELLUM. 35 

Carina moderately bowed inwards, widening gradually downwards from the apex, 
of which a small portion is filled up solid, and must have projected freely; walls 
moderately thick ; the two sides of the tectum are rather steeply inclined to each other, 
and meet in a central line, which is subcarinated with a slightly prominent ridge ; basal 
margin rectangularly pointed ; parietes nearly flat, about as wide as the tecta, in some 
specimens perpendicular, so as not to be visible when the valve is viewed from a central 
dorsal point ; in others, very steeply splayed outwards ; separated from the intra-parietes 
by a conspicuous rounded ridge, and from the tectum by a nearly equally large ridge, 
which has generally one, two, or three fine, longitudinal, raised lines on either one or both 
sides of it : in one specimen the whole surface was thus coarsely and obscurely lined. The 
intra-parietes are rather wide, extending to the basal margin of the parietes. Depth of 
valve, measured from the central crest to either inner edge, is about equal to the entire 
width, as measured from inner edge to edge. The depth compared with the width, 
though the most conspicuous character, varies a little. Inner edge of valve nearly straight. 
Length of longest specimen (in Mus. Bowerbank) 1*6 of an inch. This is the largest carina 
I have seen in any fossil cirripede. 



5. Scalpellum lineatum. Tab. II, figs. 11 and 12. 

S. superfcie totd carina lineis tenuibus, rotundatis, longitudinalibus, proximis, microcos- 
cqpicis obtectd; crista centralis costd crassiore ; costis duabus vel tribus tectum et parietes 
separantibus ; latitudine valvae circa dimidium altitudinis aquante ; intra-parietibus latius- 
culis, nulla costd conspicud a parietibus separatis ; apice solide repleto, aliquantulum libere 
prominente. 

Carina with the whole exterior surface covered with fine, rounded, longitudinal lines, 
scarcely visible to the naked eye ; with a thicker ridge on the central crest, and with two 
or three similar ones separating the tectum and parietes ; width of valve about half of 
depth; subcarinated; inter-parietes rather wide, not separated by a conspicuous ridge 
from the parietes. Apex filled up, solid, and projecting freely a little. 

Lower Chalk of Sussex, Mus. J. Morris ; Mus. J. Sowerby. 

I have seen two carina? in the collections of Mr. Morris and Mr. J. Sowerby so exactly 
like each other, and having a somewhat different aspect from S. maximum, var. sulcatum, 
to which they come nearest, that they deserve to be described, whether or not they are 
really specifically distinct. I long hesitated whether to give them a specific name, and 
have been, in some degree, influenced in doing so, from the presence of scuta and terga 
in the Lower Chalk, which indicate a distinct but closely-allied species. The scutum is in 
Mr. Morris's collection, and came in the same lot with the carina from Sussex : the tergum 



36 . FOSSIL CIRRIPEDIA. 

is in Mr. Bowerbank's collection from the Lower Chalk of Maidstone. These valves are 
marked with longitudinal raised striae more plainly than is the carina. 

Carina (fig. 12) ; moderately bowed inwards ; inner margin nearly straight ; widening 
very gradually downwards from the apex, of which a very small part is filled up solid, 
and must have projected freely ; walls rather thin. Both tecta and parietes are regularly 
striated longitudinally, with raised, hair-like, fine lines scarcely visible to the naked eye ; 
one central, and two or three on each side between the tectum and parietes, being about 
wice as large as the others, and visible to the naked eye. Tecta rather steeply inclined 
towards each other; central line sub-carinated ; basal margin rectangularly pointed; 
parietes slightly concave, about as wide as half the tectum ; steeply inclined outwards ; 
separated from the intra-parietes on each side by a slight ridge. Intra-parietes set a little 
inwards, wider in the widest part than the adjoining parietes or tecta ; extending baseward 
not as far as the basal margin of the parietes. Depth of valve measured from central crest 
to either inner edge, nearly equal to the entire width, as measured across from inner edge 
to edge. In many respects this carina is intermediate between those described under 8. 
maximum and 8. maximum, var. sulcatum ; but comes nearest to the latter : the intra- 
parietes not extending so far baseward ; and the delicately lineated exterior surface gives 
it, however, a somewhat different aspect. 

Scutum (fig. 11); this valve, from the Lower Chalk of Sussex, resembles that of 8. arcu- 
atum ; its surface is covered with raised strise, which are further apart, and less plain than 
in the typical specimens of 8. arcuatum from the Gault, but resemble those in the variety 
from the Grey Chalk of Dover. Outline trapezoidal : the baso-lateral angle is very broad, 
rounded, and protuberant ; no ridge runs from it to the apex : the basal margin projects 
very slightly close to the rostral angle, and the tergal margin is not inflected as in 8. 
arcuatum. The internal surface of the valve, along the tergal margin, is not furrowed or 
marked by lines of growth : I have no doubt that this valve is, at least, distinct from 
8. arcuatum. 

Tergum. This valve, from the Lower Chalk of Maidstone, resembles that of 8. arcuatum, 
var. from the Grey Chalk ; it is, however, slightly more elongated : it further closely 
resembles a tergum, which I have provisionally attributed to Pollicipes striatus, differing 
from it in being less elongated, and more especially in the absence of a ridge, steep on the 
carinal side, which in that species runs from the apex to the basal angle. 

Finally, I may remark, that these three valves, on the supposition that they have been 
rightly attributed to one species, indicate a form intermediate between Scalpellum maximum 
of the Upper Chalk, and 8. arcuatum of the Grey Chalk and Gault. 



SCALPELLUM. 37 



6. SCALPELLUM HASTATUM. Tab. II, fig. 13. 

S. carina intra-parietibus, intra paululum positis, instructd ; valvd totd introrsum valde 
arcuatd, margine interno non recto; margine basali acuto, lanceolato ; valvd tenui, Icevi, 
tecto transverse leniter arcuato ; parietibus a tecto vix disjunctis. 

Carina having intra-parietes set a little inwards ; whole valve much bowed inwards, 

with the inner margin not straight; basal margin sharply pointed, spear-shaped; valve 

thin, smooth ; tectum' in a transverse line flatly arched ; parietes barely separated from 

tectum. 

Grey Chalk, Dover, Mus. Brit. 

Carina smooth, narrow, furnished with intra-parietes, widening gradually from the apex 
downwards ; extremely much arcuated, so that the uppermost part, of which none, or very 
little, projected freely, is at right angles to the basal part. Not only is the dorsal surface 
considerably arcuated, but so are the inner margins, which is much more important. Basal 
margin sharply pointed, with the two edges meeting each other at about an angle of 75°. 
Roof with the two sides continuously and flatly arched ; parietes rather narrow, slightly 
concave, barely separated from the tecta : the concavity of the parietes, as seen on the basal 
margin, together with the sharpness of the central portion, makes the lower part of the 
valve spear-shaped. The intra-parietes are set a little within the parietes ; they extend 
down to about two thirds of the entire length of the carina, and not to the basal margin of 
the parietes : they are widest at about only one fourth of the entire length of the carina 
from the apex, and here they equal in width the rest of the valve. Internally the valve 
is, in the upper part, owing to the wide intra-parietes, deeply concave ; in the lower part, 
only slightly so. 

Length of carina, measured along the chord of the arch, -75 of an inch. 

Affinities. This species certainly comes very near to S. maximum ; but I think it is 
distinct, and is its representative in the Grey Chalk. I have seen only a single specimen. 
The carina differs from the former varieties and species in its smoothness, thinness, in 
the acumination of the basal margin, in its much arcuated form, and more especially (for 
this, probably, would greatly influence the outline of the terga,) in the inner margins being 
also thus arcuated. 



7. SCALPELLUM ANGUSTUM. Tab. I, fig. 2. 

Xiphidium angustum. Dixon. Geology of Suffolk, tab. xxviii, fig. 9. 

S. carina angustd, introrsum valde arcuatd ; tecto a parietibus rectangule in/lexis costd, 
(ut videtur) disjuncto ; intra-parietibus usque ad dimidium valvce pertinentibus, deinde 
oblique et abrupte iruncatis ; margine basali acute cuspidato. 



38 FOSSIL CIRRIPEDIA. 

Carina narrow, much bowed inwards ; tectum apparently separated from the rectangu- 
larly-inflected parietes by a ridge ; the intra-parietes extend down half the valve, and are 
there obliquely and abruptly truncated ; basal margin sharply pointed. 

Chalk. 

I know this species only from the plate in Mr. Dixon's work. Being well aware of 
Mr. J. de C. Sowerby's great accuracy, I cannot doubt that the intra-parietes are at 
their lower end, abruptly and obliquely truncated in the manner represented in the Plate : 
this character, with its sharply-pointed basal margin, makes me believe the species to be 
new : it comes, I imagine, nearest to S. hastatum} 



9. SCALPELLUM TRILINEATUM. Tab. I, fig. 5. 

S. carina tecto transverse leniter arcuato, subcarinato, costd centrali et costis duabus 
lateralibus, rotundatis, tumidis ; parietibus angustis leviter concavis, rectangule inflexis. 

Carina, with its tectum in a transverse line flatly arched, sub-carinated, with a central 
and two lateral, rounded protuberant ridges ; parietes narrow, slightly concave, rectan- 
gularly inflected. 

Grey Chalk, Dover, Mus. Brit., Flower. Chalk Detritus, Charing, Kent, Mus. Harris. 

Carina (fig. 5, a — d); moderately arched, narrow, gradually widening from the apex to 
the base, plainly marked by fines of growth : no part apparently projected freely. The 
tectum is flatly arched, sub-carinated, with its central crest forming a rounded protuberant 
ridge ; on each side, the tectum is bounded by similar, very slightly larger ridges, making 



1 8. SCALPELLUM QDADBICARINATUM. 

Pollicipes Q.DADRICARINATUS, Reuss, Verstein. Bohmisch. Kreideformation (1846), Tab. xlii, fig. 18. 

<S. carina intra-parietibus latis (ut videtur) instruct d ; tecto transverse piano, Icevi, costd prominente 
utrinque marginato ; margine basali abrupte truncato. 

Carina having apparently wide intra-parietes ; tectum in a transverse line, flat and smooth, bordered on 
each side by a prominent ridge ; basal margin abruptly truncated. 

Bohemia. Untern Planerkalke (Chalk-marl). 

I know this species only from an imperfect plate, but good description of a carina, in Reuss' work : it 
is an interesting form, showing in its truncated basal margin and flat tectum a still closer affinity to the 
recent S. rutilum, even than does S. fossula ; thus confirming the view I have taken of the affinities of these 
several species. 

Carina ; rather narrow, slightly bowed inwards : tectum quite flat and smooth, separated from the 
parietes by a smooth prominent ridge : parietes concave, rectangularly inflected : intra-parietes apparently 
well developed, separated from the parietes by a ridge : basal margin abruptly truncated. 



SCALPELLUM. 39 

together three ridges. The basal margin is bluntly pointed, with the two sides meeting 
each other at an angle of rather above 90°. Parietes rather narrow, rectangularly inflected, 
slightly concave : in the upper part there is no trace of intra-parietes. 

Terga (fig. 5, e — i). In Mr. Flower's collection there is a tergum, (embedded in exactly 
the same matrix,) which, from a certain degree of resemblance in outline with that of S. 
arcuatum, the nearest congener to 8. trilineatum, and from another point of resemblance 
with S.fossula, I believe belonged to this species. 1 The valve is very smooth, with obscure 
traces of fine striae radiating from the umbo ; nearly flat ; pointed oval, but with the scutal 
side much more protuberant than the carinal. Apex much acuminated, curled forwards ; 
carinal margin much and regularly bowed from the upper to the basal point, which latter 
is blunt and square : from it to the apex there runs, in a curved line, nearly parallel to the 
carinal margin, a barely perceptible broad ridge. Occludent margin curved up towards 
the umbo, short compared to the scutal margin ; parallel to it there runs a very wide and 
very shallow depression. Scutal margin, with a portion corresponding with the above 
depression, forming rather more than a third of the margin, not projecting so much as the 
lower two thirds, and separated from this lower part by a slight bend, probably marking 
the spot to which the apex of the upper latera extended. 

Affinities. The carina obviously most resembles that of S. fossula and arcuatum ; it 
differs plainly from both, in having a centra] rounded ridge : in the two well-developed 
boundary ridges of the tectum it comes nearest to the cretaceous iS. fossula ; but in the 
absence of the intra-parietes (and this I conceive is a more important character), it comes 
nearest to the S. arcuatum, from which, however, it can be at once distinguished by the 
absence of the longitudinal stria?. The tergum above described, which I believe belonged 
to this species, in the form of the scutal margin, comes nearest to that of S. fossula, 
though in general shape perhaps nearer to S. arcuatum. In /S.fossula the carina has intra- 
parietes, which are closely adjusted to the straight carinal margins of the terga : in S. 
trilineatum the intra-parietes are absent, but in their place the carinal margins of the two 
terga are themselves highly protuberant, so that in these two species, although the upper 
parts of the carina? and terga are separately of very different shapes, they give, when com- 
bined together, a similar general outline. 



10. SCALPELLUM SIMPLEX. Tab. I, fig. 9, 

8. carina leevi; parietibus angustissimis, rectangule inflexis ; tecto subcarinato, trans- 
verse mediocriter arcuato ; margine basalt rectangule acuminato. 

1 It must however be added that the terga, at present unknown, of S. hastatum, a species occurring in 
the Grey Chalk of Dover, would probably have the same outline, and almost certainly would have a very 
smooth surface. 



40 FOSSIL CIRRIPEDIA. 

Carina smooth ; parietes extremely narrow, rectangularly inflected ; tectum sub-cari- 
nated, in a transverse line moderately arched ; basal margin rectangularly pointed. 

Lower Greensand, Maidstone. Mus. J. Morris. 

I know this species only from a single carina, which is chiefly characterised by its 
simplicity : it is, I think, certainly distinct from all the others. In the sides of the carina 
being simple, that is in not being divided by a ridge into parietes and intra-parietes, it 
comes nearest to 8. arcuatum and trilineatum, from the former of which it is readily distin- 
guished by its smoothness ; and from 8. trilineatum by the absence of the three ridges. 
This species possesses some interest, as being the oldest cirripede, which I have ventured 
to attribute to the genus Scalpellum. Carina moderately tapering, slightly bowed towards 
the terga; sub-carinated, but with the central ridge smooth; transversely moderately 
arched; basal margin rectangularly pointed; the whole surface is smooth. Parietes 
extremely narrow, rectangularly inflected, set inwards, not extending down to the basal 
margin, with the lines of growth almost parallel to the inner edges of the valve. 



11. Scalpellum arcuatum. Tab. I, fig. 7. 

S. valvarum lineis angustis elevatis ab apice radiantibus : carina tecto transverse leniter 
arcuato, et parietibus rectangule i?iflexis, leniter concavis, lavibus. 

Valves with narrow elevated lines radiating from their apices. Carina with its tectum 
in a transverse line flatly arched, and with the parietes rectangularly inflected, slightly 
concave, smooth. 

Gault, Folkstone, Mus. Bowerbank, J. Sowerby, Flower. Var. in Grey Chalk, Dover, Mus. Brit. 
Planer (Chalk-marl) near Hildesheim. Mus. Roemer. 

I have ranked this species under Scalpellum instead of Pollicipes, from the somewhat 
greater resemblance of its scuta and carina with the fossil species of Scalpellum, than 
with any known Pollicipes ; though in some respects it appears rather intermediate. 
This species appears to come nearest to the Pollicipes radiatus of J. de C. Sowerby in 
* Geol. Trans./ vol. iv, 2d Series, PL XI, fig. 6, but besides that that species comes from 
the Lower greensand, the lower angle of its tergum is much more pointed; the upper 
figure of the two there given appears to be something wholly different. From the state of 
the specimens, I believe that the three following valves, all in Mr. Bowerbank's collection, 
belonged to the same species. 

General Appearance. Carina, scuta, and terga plainly marked with prominent, very 
narrow, straight ridges, radiating from their apices; the interspaces between these ridges are 



SCALPELLUM. 41 

three or four times as wide as the ridges themselves ; the lines of growth are very fine and 
narrow. 

Carina (fig. 7, a, b, g) \ narrow, considerably arched : tectum flatly arched, obscurely 
subcarinated : parietes rectangularly inflected, somewhat concave, and not longitudinally 
ridged, like the tectum, about two thirds as wide as half the tectum : basal margin bluntly 
pointed, the two edges meeting each other at rather above a right-angle ; a trace of a 
rounded ridge separates the tectum and parietes; in the upper part of the carina there 
is no trace of intra-parietes, therefore the section of the upper half of the carina is only 
four- sided, see fig. 7, g. 

Scutum (fig. 7,/) ; moderately convex, with the apex acuminated : lateral margin nearly 
parallel to the slightly arched occludent margin, and at right angles to the straight 
basal margin ; a distinct ridge runs from the apex to the baso-lateral angle, which is 
distinctly prominent and rather sharp. The valve, above a line running from the apex 
to the tergo-lateral angle, is inflected; and the narrow portion thus inflected, which 
cannot be seen when the valve is viewed from above, is destitute of the longitudinal 
ridges. 

In a specimen from the Grey Chalk of Dover, in which the internal surface was visible, 
there was, above the well-marked depression for the adductor muscle, a prominent, central, 
slightly oblique ridge, with the inner occludent edge of the valve widened and slightly 
hollowed out on the one side, and with a trace of a furrow on the other or tergal side. 

Terga (fig. 7, c, d) ; flat, oval, with the scutal angle rather protuberant ; basal angle not 
sharply pointed, from it to the apex there runs an obscure furrow, which furrow in the lower 
part of the valve is central, but higher up is situated at about one third of the width of 
the widest part of the valve from the carinal margin ; in the lower part of the valve, the 
lines of growth (and consequently the margins of the valve) make with this furrow, equal 
angles on its opposite sides. The valve is slightly depressed, parallel to the occludent 
margin. A small portion of the apex projected freely ; internally, in the upper part, 
rather nearer to the occludent than to the carinal margin, the valve is prominent, and 
this part is marked with two or three little ridges (c) ending abruptly downwards. 

Size of largest specimen, — length of carina, # 85 of an inch; of scutum, from the 
apex to the basal margin, rather above '6; of terga, '55. I do not, however, know 
that these valves belonged to the same individual. 

Variety. In the British Museum there is a scutum/and in Mr. Flower's collection 
there is a tergum, both from the Grey Chalk of Dover, which are most closely allied to, 
if not identical with, the above valve. The raised striae on both are rather further apart 
and are less prominent. In all the other characters the scutum is identical. The tergum 
differs in its carinal margin, being rather more angularly bent, and in there being no furrow 
running from the apex to the basal angle ; but these differences are trifling and insufficient 
for distinguishing a species. Amongst some specimens most kindly sent me by Roemer, 
there is a tergum from the Planer of Sarstedt (Chalk-marl), which is identical with this. 

f 



42 FOSSIL CIRRIPEDIA. 

Affinities. This species is related to S. trilineatum, simplex, and solidulum, in the 
absence of intra-parietes ; in the terga it comes closest to the latter species. 1 

1 12. SCALPELLUM SOLIDULUM. Tab. I, fig. 8. 

Pollicipes solidulus. Steenstrup in Kroyer's Tidsskrift, b. ii (1839), pi. v, fig. 14 et 14*. 
— undulatus. Id. Id. Id. fig. 6. 

S. valvarum lineis latiusculis elevatis ab apiee radiantibus. Carince parte superiori liberh prominente, 
et crista centrali, internd, longitudinali instructd. 

Valves with rather wide elevated lines radiating from their apices. Carina, with the upper part freely 
projecting, and internally urnished with a central prominent, longitudinal crest. 

Scania (Kjuge). Mus. Univers. Copenhagen. 

Professor Steenstrup has described under this name some carinse, in so worn a condition, that I confess 
that I thought it quite impossible to characterise them ; and under the name of P. undulatus, some well- 
preserved terga. Quite lately, M. Angelin has sent to Professor Steenstrup, from Kjuge in Scania, several 
of the same carinse in a much better condition, a scutum, and some broken terga of P. undulatus, which, 
from the similarity of their longitudinally striated surfaces, M. Angelin believes belonged to the same 
species : I quite concur in the probability of this view. The better state of the carinse proves the sagacity 
of Professor Steenstrup, in considering his worn specimens indicative of a distinct species. Had I seen 
these carinse alone, I should have much hesitated in considering them as belonging to a Scalpellum : for 
they differ considerably from the same valve in all other species ; the parietes, or rather the part answering 
to the parietes, being here so much inflected, that they fill up and render solid the upper part of the valve ; 
but the scutum undoubtedly belonged to a Scalpellum, and the terga closely resemble the same valve in the 
£. areuatum. 

Carina (fig. 8, b, c, d) ; narrow, elongated, strong and solid ; moderately bowed inwards ; basal margin 
rectangularly pointed ; surface covered with rather broad slight ribs, central one being apparently (for the 
best specimens are much worn) twice as broad as any of the others. In a transverse line, the tectum is 
considerably arched in the upper part of the valve, and only slightly arched in the lower part. A considerable 
length of the upper part must have projected freely ; this portion being filled up solid, and having a central, 
largely prominent crest or ridge : it appears, for the specimens are in a much worn condition, as if the 
ridge had been formed by the inflection of the parietes on each side, and their perfect junction. In the 
peculiar and almost monstrous variety of S. maximum, called var. cylindraceum, we have nearly the same 
structure ; a cleft, however, being left, marking the line of junction of the opposite parietes. In general 
appearance and proportions, this carina comes nearest to those of Scalpellum ; but in the peculiar modi- 
fication of the parietes (if they can be so called) into a central crest, and in the apparent (from worn state) 
absence of any distinct ridge separating the tectum and parietes, the valve departs from the general 
description of the carina in Scalpellum. 

Scutum ; of this valve, which undoubtedly belonged to a Scalpellum, there is one entire specimen, 
but with the angles so much rounded, that I can point out no distinguishing character from the same 
valve in 8. areuatum (fig. 7, g), of which a figure has been given, except that the longitudinal ridges 
are proportionally broader and further apart. The ridges closely resemble those on the above-described 
carina. 

Terga (fig. 8, a) ; sub-triangular, flat, strong, and thick, with moderately wide, not quite straight ridges, 
radiating from the apex : the interspaces betwen the ridges are three or four times as wide as the ridges 
themselves ; valve very slightly depressed, parallel to the occludent margin. A slight ridge, connecting the 
sharp basal apex, runs quite close to the carinal margin, even in the lower part of the valve: in 



SCALPELLUM. 43 



13. SCALPELLUM TUBERCULATUM. Tab. I, fig. 10. 

8. valvarum lineis tenuibus, tuberculatis, elevatis, ab apice radiantibus : carina tecto 
transverse leniter arcuato, et parietibus striatis; scuti umbone prope in medio marginis 
occludentis posito, costis duobus ab umbone ad angulum basi-lateralem, et ad basalis marginis 
medium decurrentibus. 

Valves, with fine, tuberculated, elevated lines, radiating from their apices : carina, with 
the roof in a transverse line, gently arched, and with the parietes striated : scutum, with 
the umbo placed nearly in the middle of the occludent margin, with two ridges running 
from the umbo to the baso-lateral angle and to the middle of the basal margin. 

Chalk Detritus. Charing, Kent. Mus. Harris. 

Through the kindness of Mr. Harris, I have examined several valves, which I believe 
to belong to the same species : the specimens were found in the chalk detritus, and, 
therefore, may have come from the Upper or Lower Chalk or Chalk-marl ; but more 
probably from the Upper Chalk. With respect to the scuta and terga I have scarcely any 
doubt, from certain peculiarities, that they belonged to the same species ; but with regard 
to the most important valve, the carina, I cannot feel quite so certain : when the latter is 
so held, that the parietes are not visible, it has a very close general resemblance to the same 
valve in Pollicipes rigidus. In the carina, the present species comes closer to JS. arcuatum 
than to any other species ; in the other valves, especially in the singular scuta, it departs 
widely from that and all other known fossil forms, with the exception of 8. (?) cretcs, of 
Denmark. All the specimens which I have seen are small; the carina being -2 long, and 
the terga less than '15 of an inch in length, in the largest specimens. 

consequence of this, the lines of growth make a different angle, on the opposite sides, with this ridge : 
as the valve has heen somewhat worn, it is possible that the carinal margin may have been more abraded 
than is apparent. Internally, it is seen that a considerable portion of the upper part of the valve projected 
freely ; beneath this, the inner surface is slightly convex, but smooth, and though the shell has been much 
worn, I doubt whether there ever existed ridges, as on the internal surface of the upper part of the terga 
in S. arcuatum, to which valve this presents a close general resemblance. Length of tergum (when 
perfect), 1-2 of an inch. 

Carinal Latus (fig. 8, e, f) ; amongst the fossils from Kopinge (at which place the same species are found 
as at Kjuge), there is a valve, which I believe to be a carinal latus of a Scalpellum, and which, from its 
longitudinal ridges, more probably belonged to the present than to any other species : from its peculiarity 
it is in any case worthy of description. In form it is a segment, somewhat less than a quarter, of a circle ; 
of this segment, nearly half (I believe the upper half) has its end or circumferential margin much 
hollowed out, and its surface smooth : the other half has its periodical growth-ridges very prominent, 
and these are crossed by a few slight longitudinal ridges. One of the lateral sides (the upper, I believe,) 
is reflexed so as to form a prominent ledge ; the other side is slightly inflected. 



44 FOSSIL CIRRIPEDIA. 

The valves all have their surfaces plainly ribbed longitudinally ; the ribs are narrow, 
and as they cross each zone of periodical growth they are tuberculated. 

Carina (fig. 10, b, c); narrow, tapering, little bowed inwards ; tectum in a transverse line, 
steeply arched, not carinated ; basal margin bluntly pointed ; in very young specimens, 
however, it is evident from the lines of growth, that the basal margin must have been 
rounded; the parietes are inflected, and rather narrow, being barely half the width of 
half the tectum; they are plainly marked by parallel lines of growth; internally the valve 
is rather deeply concave ; no part of the apex projected freely. 

Scuta (fig. 1 0, e) ; umbo of growth on the occludent margin, at about one third of the 
entire length of the valve from the apex ; somewhat convex ; four-sided, the margins 
consisting of the lateral, which is considerably longer than the other sides ; the basal 
which forms nearly a right angle with the lower half of the occludent margin ; and 
of an upper and lower occludent margin, meeting each other at about an angle of 135° : 
the margin which I have here called the upper occludent, homologically corresponds with 
the tergal margin of the other cretaceous species, and with the upper, nearly straight, 
portion of the occludent margin in the tertiary S. magnum and the recent JS. vulgare, — 
a fact which has been mentioned under the head of Scalpellurn. The edge of the upper 
occludent margin forms a strongly prominent ridge, with its apex forming a slight pro- 
jection ; a second less prominent ridge runs from the umbo to the baso-lateral angle, and a 
third faint ridge runs from the umbo to a point in the basal margin, nearer to the rostral 
than to the baso-lateral angle. Internally there is a rather deep hollow for the adductor 
muscle; along the under surface of the upper occludent margin there is a slightly 
prominent ridge, bordered by two slight depressions. 

Terga (fig. 10, a); flat, elongated diamond-shape; close and parallel to the occludent 
margin there is a narrow, very prominent ridge or plait, the end of which forms a slight 
projection; a straight ridge runs from the apex to the sharp basal angle; the scutal and 
lower carinal margins are of equal length, and longer than the occludent and upper carinal 
margins, which latter are equal, and meet at an angle very slightly less than a rectangle. 
On the under surface there is a slight depression and ridge, close and parallel to the 
occludent margin. I have no doubt that the ridge along the upper occludent margin of 
the scuta, and that on the occludent margin of the terga, together with their projecting 
points, are related to each other, owing to the close contact of these valves. 1 

1 14. ScALPELLrM SEMIPOKCATTM. Tab. I, fig. 6. 

S. carina ignotd : scirti costis duobus ab umbone ad unguium basi-lateralem et ad marginis basalts 
medium decurrentibus : superficie inter hanc costam et marginem occludentem lineis tenuibus, longitudinalibus, 
elevatis instructd. 

Carina unknown : scutum, with two ridges running from the umbo to the baso-lateral angle, and to 



SCALPELLUM. 45 



15. SCALPELLUM (?) CRET.E. Tab. I, fig. 11. 

Anatifera cret^i. Steenstrup. Kroyer's Tidsskrift, 1837 et 1839, b. ii, pi. v, 
figs. 1, 2, 3. 

S. valvis Iambus tenuissimis : scuti umbone prope medium marginis occludentis posito ; 
eostis tribics obscuris ab umbone ad angulos tergo-lateralem et basi-lateralem, et ad medium 
marginis basalis decurrentibus : carina apice et margine basali acutis; distincti parietes absunt. 

Valves smooth, extremely thin : scutum with the umbo placed nearly in the middle of 
the occludent margin, with three obscure ridges running from the umbo to the tergo-lateral 
and baso-lateral angles, and to the middle of the basal margin : carina with the apex and 
basal margin sharply pointed ; without distinct parietes. 

White Chalk, Denmark, Mus. Univers., Copenhagen. Chalk Detritus, Charing, Kent (?), Mus. Harris. 

Preliminary Remarks. I owe to the kindness of Professor Steenstrup, as in so many 
former instances, an examination of several specimens of this fossil, which is of interest, as 
being extremely common and characteristic of the white chalk of Denmark. Amongst 
the numerous minute specimens from the chalk detritus of Charing in Kent, sent me by 
Mr. Harris, there are some carinas so similar that I have ventured, with doubt, to rank this 
as a British species ; the carina, however, in this species, are far from characteristic. I have 
felt much hesitation in admitting this species in the genus Scalpellum : Professor Steenstrup 
was originally inclined to believe that the capitulum was formed of only five valves ; 
could this be proved, the species would very naturally rank with a small recent one from 
the Island of Madeira, which, owing to the upward growth of the scuta, and to certain 
peculiarities in the animal's body, I have felt myself compelled to raise to the rank of a 
genus, under the name of Oxynaspis. But as the valves of S. (?) cretce have never been found 
united, and as the main ones are very small, fragile, and generally in a broken condition, 
the small lower ones might easily be overlooked. I have seen, indeed, in two instances, 

the middle of the basal margin ; the surface between the latter ridge and the occludent margin covered 
with fine longitudinal elevated lines. 

Scania (Kopinge). Mus. Univers., Copenhagen. 

I have in this one instance departed from my rule of never naming any other valve, except the carina 
in the genus Scalpellum ; but the scutum here to be described almost certainly belongs to this genus, and 
is interesting in connection with the homologous valves in <S. tuberculatum and S. (?) cretce, to which 
species it is apparently allied, but yet differs greatly from them in the umbo being seated at the upper- 
most point of the valve. 

Scutum, moderately elongated, slightly convex ; a narrow, prominent, well-defined ridge runs from the 
apex to the baso-lateral angle, at which point it forms a narrow projection : a second ridge, not quite 
so prominent, runs from the apex to the basal margin, to a point rather nearer to the baso-lateral than to 
the rostral angle. That part of the valve between this second ridge and the occludent margin has four or 
five faint longitudinal ridges, whereas the rest of the valve is smooth. Internally there is a deep depression 
for the adductor muscle, above which the surface is simply concave up to the apex. 



46 FOSSIL CIRPJPEDIA. 

what appeared to be upper latera, but as I could not remove them so as to examine their 
under sides, I am far from sure that they were not broken, angular portions of scuta. If 
we look to the character of the separate valves, there is a striking and important resem- 
blance between the scuta of S. creta and tuberculatum, in the umbo being seated in a 
nearly middle point of the occludent margin, and likewise in the two ridges running from 
the umbo to the baso-lateral angle, and to a central point of the basal margin ; in which 
latter character of the ridges, this species also agrees with S. semiporcatum. These facts 
have determined me, provisionally, to rank the present species under Scalpellum. But on 
the other hand, if we look to the carina, which, according to our rule, is considered the 
characteristic valve in this genus, it rather resembles the homologous valve in Pollicipes ; 
for the carina has not any parieties separated from the tectum by a distinct ridge. The 
terga seldom afford any serviceable generic characters ; but as far as they go, they also 
rather resemble the terga in Pollicipes than in Scalpellum. Hence, it is obvious, that the 
generic position of S. (?) cretce is at present very uncertain. 

Valves small, smooth, extremely thin and brittle. 

Scutum (fig. 11, c); trapezoidal, with the upper part of the valve produced into a sharp 
point, and with the rostral angle slightly and obliquely cut off. Umbo seated at a little 
above the middle of the occludent margin, which is straight. The tergal margin is longer 
than the lateral margin : the basal margin (on the carinal side of the truncated rostral end) 
forms a right angle both with the lateral and occludent margins. Valve somewhat convex 
near to the umbo, whence three obscure ridges radiate, — one to the angle between the tergal 
and lateral margins ; a second to the baso-lateral angle, and a third to the bend in the basal 
margin ; these ridges, however, seem to vary in strength, and in the largest specimens 
could hardly be distinguished : in most of the specimens, the narrow portion of the valve 
which ends in the truncated rostral angle, is a little inflected. The lines of growth follow 
the basal and tergo-lateral margins, and can be traced just bending round the sharp 
apex, so that a very narrow ledge is added along the upper part of the occludent margin. 

Tergum (fig. 21, a); sub-rhomboidal, nearly flat: the carinal margin consists of an 
upper larger portion, and of a lower, shorter portion : the occludent and scutal margins 
are nearly equal in length. The apex is a little curled towards the scuta, and is sharp ; 
basal angle bluntly pointed. A faint curved ridge runs from the apex to the basal angle, 
at about one fourth of the entire width of the valve from the carinal margin. 

The Carina (fig. 11, h) widens rapidly downwards from the extremely sharp apex; 
basal margin spear-shaped, sharply pointed, the two edges meeting each other at about an 
angle of 75°; exterior surface sub-carinated; in a transverse line the valve is slightly arched, 
and longitudinally, very slightly bowed inwards : with a lens, traces of longitudinal striae 
are visible. 

Dimensions. The species seems to have been always small : the largest scutum and 
tergum were each about a quarter of an inch in length. Probably the individuals were 
attached in groups to corallines at the bottom of the cretaceous sea. 



POLLICIPES. 47 



Genus — Pollicipes. 

Pollicipes. Leach. Journal de Physique, torn, lxxxv, Julius, 1817. 1 
Lepas. Linn. Systema Naturae, 1767. 
Anatifa. Brugiere. Encyclop. Method, (des Vers), 1792. 
Mitella. Oken. Lehrbuch der Naturgesch., 1815. 

Ramphidiona. Schumacher. Essai d'un Nouveau Syst. &c, 1817 (ante Julium). 
Polylepas. De Blainville. Diet, des Sc. Nat., 1824. 

Capitultjm (secundum Klein). J. E. Gray. Annals of Philos., torn, x, 2d series, 

Aug. 1825. 

Valvce ab octodium usque ad centum et amplius. Lateribus verticelli inferioris multis ; 
lineis incrementi deorsum ordinatis. Subrostrum semper adest. Pedunculus squamiferus. 

CHARACTERES VALVARTJM IN SPECIEBUS FOSSILIBUS. 

Carina ab apice ad marginem basalem multum dilatata ; apex plerumque libere pro- 
minens ; parietes a tecto non distincte separati ; linece incrementi parietum parum obliques. 
Scuta plerumque subsolida, convexa, subtrigonalia, margine tergo -later ali plus minusve 
eminente, sed non angulo in margines duos discreto. 

f . Scuta, aid Icevia aid lineis tenuibus incrementi solum notata. 

A. Scuta, costd ab apice ad centrum marginis basalts non decurrente. 

B. Scuta, costd, nonnunquam subobsoletd, ab apice ad centrum marginis basalis 
decurrente. 

tf. Scuta, aid longitudinaliter aut transverse (i. e. secundum lineas incrementi) costata. 

Valves from eighteen to above one hundred in number. Latera of the lower whorl 
numerous, with their lines of growth directed downwards. Sub-rostrum always present. 
Peduncle squamiferous. 

CHARACTERS OF THE VALVES IN FOSSIL SPECIES. 

Carina ; widening considerably from the apex, which projects freely, to the basal 
margin ; parietes not distinctly separated from the tectum ; lines of growth on the 
parietes but little oblique. Scuta generally somewhat massive, convex, sub-trigonal, with 

1 This is one of the rare cases in which, after much deliberation and with the advice of several 
distinguished naturalists, I have departed from the rules of the British Association ; for it will be seen that 
Mitella of Oken, and Ramphidiona of Schumacher, are both prior to Pollicipes of Leach ; yet as the latter 
name is universally adopted throughout Europe and North America, and has been extensively used in 
geological works, it has appeared to me to be as useless as hopeless to attempt any change. It may be 
observed that the genus Pollicipes was originally proposed by Sir John Hill (' History of Animals,' vol. iii, 
p. 170), in 1752, but as this was before the discovery of the binomial system, by the Rules it is absolutely 
excluded as of any authority. In my opinion, under all these circumstances, it would be mere pedantry to 
go back to Oken's ' Lehrbuch der Naturgesch.' for the name Mitella, — a work little known, and displaying 
entire ignorance regarding the Cirripedia. 



48 FOSSIL CIRRIPEDIA. 

the tergo-lateral margin more or less protuberant, but not divided by an angle into two 
distinct margins. 

t Scuta smooth, or marked only with fine lines of growth. 

A. Scuta without any ridge proceeding from the apex to a nearly middle point of the 
basal margin. 

B. Scuta with a ridge, sometimes faint, proceeding from the apex to a nearly middle 
point of the basal margin. 

ff . Scuta either longitudinally or transversely (that is in the direction of the lines of 
growth) ridged. 

As with Scalpellum, the first of the above two paragraphs contains the true generic 
description, as applicable to recent and fossil species ; the second paragraph has been drawn 
up as an aid in classifying separated valves. This, the most ancient genus of the Lepadidse, 
seems also to be the stem of the genealogical tree; for Pollicipes leads, with hardly a 
break, by some of its species into Scalpellum villosum ; and Scalpellum leads by Oxynaspis 
into Lepas and the allied genera : Pollicipes mitetta, moreover, is nearer allied to the 
Sessile Cirripedes than is any other Pedunculated cirripede, except, perhaps, Lithotrya, which 
is also closely connected with Pollicipes. The six recent species of Pollicipes might be 
divided into three sub-genera : one containing the P. mitella ; a second, P. cornucopia, 
elegants and polymerus; and the third, P. spinosus and serta (nov. spec.) Of the fossil species 
some, as P. carinatus, dorsafus, validus, &c, are related to the first section ; others, as 
P. reflexus and concinnus, to the second section ; and lastly, others, as P.glaber and unguis, 
perhaps form a distinct section, though more related to P. mitella than to other recent 
species. As, however, most species are known by only a few of their valves, it is scarcely 
possible to speak with certainty regarding their finer affinities. 

Description : as in the case of Scalpellum, the following remarks are confined to the fossil 
species of the genus. In all full-grown recent species the number of valves in the capitulum is 
very large : this seems to have been the case with the Oolitic P. concinnus, and probably with 
most other species, but whether with all may be doubted ; from the size of the carinal 
latera of the lower whorl in P. unguis, I suspect that the total number of its valves cannot 
have been great. The valves are either smooth or plainly marked by the lines of growth, 
or they rarely have longitudinal ridges, or transverse ridges corresponding to each 
periodical zone of growth : no recent Pollicipes has a surface of this latter kind. The 
valves in Pollicipes are often strong and massive, with their apices projecting freely from 
the capitulum. 

Scuta generally three-sided, but sometimes, from either the baso-lateral or rostral 
angles being truncated, there is an additional lower side. The tergo-lateral margin is 
either straight or generally more or less convex, but it is never (as far as I know) divided 
into two distinct margins, as is always the case with Scalpellum owing to the abrupt 
ending of the upturned lines of growth. The basal margin is either straight or formed 



POLLICIPES. 49 

of two lines meeting each other at a wide angle, or somewhat irregular. The angle which 
this basal margin makes with the occludent margin varies much. The occludent margin is 
slightly arched, and is sometimes exteriorly strengthened by a ledge or rim. A prominent 
ridge runs in several species from the apex of the valve to the baso-lateral angle; 
and in another set of species there is a second obscurer- ridge running to a nearly middle 
point of the basal margin : in this latter set, the two ridges no doubt mark the extent to 
which the rostrum and upper latera overlapped the scutum. Internally there is almost 
always a deep pit for the adductor scutorum muscle : the upper part of the valve generally 
projects freely, and is internally marked by lines of growth ; sometimes there is a furrow 
along the upper part either of the occludent or the tergal margin ; in the latter case the 
furrow seems to receive the scuto-occludent angle of the adjoining tergum, and thus locks 
the two valves together, as in the recent P. mitella. In two species the occludent margin 
at the rostral angle is internally produced downwards into a depending tooth or projection. 

Terga : nearly flat, rhomboidal or sub-rhomboidal ; a line formed by the converging zones 
of growth, or a ridge, sometimes steep only on the carinal side, sometimes steep on both 
sides, runs from the apex to the basal angle. The basal angle is sometimes truncated. 

Carina : is either bowed inwards or is straight : it widens from the apex down- 
wards more rapidly than in Scalpellum ; generally a considerable upper portion projects 
freely ; this upper portion is always much less concave than the lower part : it is sometimes 
filled up flat, and sometimes has even a central prominent crest ; the basal margin is either 
bluntly pointed, rounded, or truncated ; the parietes are generally more or less inflected, 
but they are not separated by any defined ridge or angle from the roof or tectum ; the 
lines of growth on the parietes are transverse, or generally only slightly oblique. These 
characters will, I believe, in nearly all cases serve to distinguish the carina of a Pollicipes 
from that of a Scalpellum. 

Sub-carina : I know of the existence of this valve only in P. concinnus, but I cannot 
doubt that it existed in all, or nearly all, the species. I have sometimes suspected that it 
might possibly have been absent in P. unguis and glaber, in which the carinal latera are so 
large. 

Rostrum and sub-rostrum : as these valves occur in P. unguis, I have little doubt that 
they are universal ; they are apparently present in P. concinnus ; the rostrum always resem- 
bles the carina, but is shorter and proportionally broader ; a larger proportion, also, seems 
always to have projected freely, caused no doubt by the more abrupt flexure of this 
end of the capitulum : this latter character is the most certain one by which the rostrum 
may be distinguished from the carina. The sub-rostrum in P. unguis resembles the rostrum, 
but is smaller, and exteriorly is not carinated. 

Upper latera .- I know these only in P. unguis and glaber, in which they consist of a 
flat triangular plate, and in P. concinnus, in which they seem to be diamond-shaped. Lower 
latera, these in P. concinnus also seem to be diamond-shaped, as in P. cornucopia ; in 
P. unguis and glaber the apices of these little valves do not project freely, and they 

g 



50 FOSSIL CIRMPEDIA. 

have a different appearance from their homologues in any recent species : they are 
trigonal, with their basal margin rounded and one end produced, to which end a narrow 
well defined ridge runs obliquely from the apex of the valve. 

The peduncle is known only in P. concinnus ; in this species it is covered with minute 
quadrangular calcified scales. 

f. Scuta, aut Icevia aut lineis tenuibus increments solum notata. 
A. Scuta, costd ab apice ad centrum marginis basalts non decurrente. 
1. Pollicipes Concinnus. Plate III, fig. 1. 

Pollicipes concinnus. J. Morris. Annals of Nat. Hist., vol. xv, 1845, p. 30, 

pi. vi, fig. 1, et Mineral Conch., pi. 647, fig. I. 

P. scutis pane quadratis, margine basali prope rostrum subconcavo, segmento tergo- 
laterali, e lineis incrementi tit videtur reflexis formato, lato, rotundato et prominente : tergis 
latis, pane quadratis : carina margine basali, tit videtur acuto. 

Scuta, almost square, with the basal margin near the rostrum a little hollowed out ; 

tergo-lateral slip, apparently formed by upturned lines of growth, broad, rounded, and 

protuberant. Terga broad, almost square. Carina, with the basal margin apparently 

pointed. 

Oxford Clay, Middle Oolite, attached to an Ammonite. Mus. Pearce. 

Although to my great regret the state of Mr. Pearce's health has prevented him 
allowing me to examine the specimens in his possession, yet I have thought it advisable to 
commence the genus with this species, as it is in a far better state of preservation than any 
other specimen hitherto discovered. We gain by a single glance the knowledge that at so 
remote a period as the Middle oolite a true Pollicipes existed. In no other instance that I 
have heard of, has the peduncle been perfectly preserved. Mr. Morris first named and briefly 
described this interesting species ; subsequently Mr. James Sowerby has given enlarged 
drawings (without any description) of it in the ' Mineral Conchology ;' and it is from these 
figures that T have drawn up my specific description, which, from this cause, is necessarily 
imperfect. The figures in this volume are copied from those in the ' Mineral Conchology,' 
which I may remark have evidently been executed with great care, and Mr. Sowerby's 
accuracy of observation is universally well known. The peduncle is several times longer 
than the capitulum -. Mr. Morris describes the scales on the peduncle as being small, closely 
pressed together, somewhat quadrate in form, and each regularly marked by a transverse 
carinated ridge ; this latter character I do not understand. The rostrum is not clearly 
figured by Mr. Sowerby, but I believe that I can see evidence of its existence. From 
these materials it would appear that the P. concinnus is more nearly related to the recent 
P. cornucopia, and its two nearest congeners, than to the other species of the genus. 



POLLICIPES. 51 

2. Pollicipes ooliticus. Tab. Ill, fig. 2. 

Pollicipes ooliticus. Buckman. Outline of the Geology of Cheltenham, by Sir R. 
Murchison, new edit, by James Buckman and H. Strickland, 1845, Tab. iii, fig. 7. 

P. scutis triangulis ; sujjerjicie undulatd ; margine basali rectangule ad marginem rectum 
tergo-lateralem posito ) segmentum tergo-laterale a lineis refiexis incrementi formatum deest. 
Carina pane redd, semicylhidricd, margine basali quadrato. 

Scuta triangular ; surface undulatoiy ; basal margin at right angles to the straight 
tergo-lateral margin ; there is no tergo-lateral segment formed by upturned lines of growth. 
Carina nearly straight, semicylindrical, with the basal margin square. 

Stonesfield Slate, Lower Oolite : Eyeford. Mus. Buckman, and Geolog. Soc. 

My materials consist of several scuta, terga, and carina?, kindly lent me by Professor 
Buckman, and of another set (which includes the rostrum) presented by him to the 
Geological Society of London. 

Valves : these have a smooth surface, but are undulatory in the direction of the lines of 
growth ; at the cessation, apparently, of each zone of growth, there was a tendency to form 
a projecting ridge or plait, as takes place in a far more marked manner in some of the 
cretaceous species, namely, P. elegans and fallax. There are also excessively fine, longitu- 
dinal striae, which can be seen only when the valves are held in particular lights ; these 
seem to have been formed by the so-called epidermis, which we know in the recent P. 
mitella is longitudinally and finely ribbed. Scuta (fig. 2, c) but slightly convex ; triangular ; 
basal margin straight, forming a right angle with the tergo-lateral margin, and rather less 
than a right angle with the slightly arched occludent margin ; the tergo-lateral margin is 
straight, and not at all protuberant : in the figure the left hand is, as usual, the occludent 
margin; I mention this because the valve has a reversed appearance, owing to the unusually 
small angle which the occludent makes with the basal margin. Terga (fig. 2, d) rhom- 
boidal, slightly convex, with a rounded ridge, which is central, running from the apex 
to the broad, rounded basal angle ; the upper carinal and occludent margins stand at 
right angles to each other, and are short compared to the scutal and lower carinal margins ; 
there is no trace of a depression parallel to the occludent margin. Carina (fig. 2, a, b) 
elongated, triangular ; scarcely at all bowed inwards ; not even <s?^-carinated ; basal margin 
rounded, not at all protuberant. The Bostrum differs from the carina only in its greater 
breadth compared to its length. 

Dimensions. The largest scutum is *6 long, but as there is a broken tergum about 11 
long, no doubt the species attained a rather large size ; the longest carina is "7 in length. 

Diagnostic characters. This species is best characterised by the straightness of the whole 
tergo-lateral and of the basal margin of the scuta; by the ridge being central on the terga; 
by the carina not being carinated ; and by the sinuous state of the surface of the valves, 
intermediate between the smooth species and those with distinct ridges parallel to the zones 



52 FOSSIL CIRRIPEDIA. 

of growth. The remarkable straightness of the tergo-lateral margin of the scuta is like that 
in the recent P. spinosus and serta, and in Scalpellum villosum, in all which species, I may 
observe, the scuta and terga are separated by an interspace of membrane ; in these three 
recent species, however, the basal margin is considerably protuberant. The present species 
differs apparently from the P. concinnus of the Oxford clay, in the basal and tergo-lateral 
margins of its scuta being straight ; in the greater proportional length of the scutal and 1 

1 3. Pollicipes Nilssonii. Tab. Ill, fig. 11. 

Pollicipes Nilssonii. Steenstrup. Kroyer, Naturhist. Tidsskrift, 1839, pi. v, figs. 20 — 23. 

P. scutis triangulis, plants : margine basalt cum margine occludente angulum pcene rectum, cum margme 
recto tergo-laterali, angulum aliquanto minor em formante. Deest segmentum tergo-laterale, lineis incrementi 
refiexis formatum. Carind introrsus admodum arcuatd, crassd ; marginis basalts mucrone obtuso. 

Scuta triangular, flat ; basal margin forming nearly a rectangle with the occludent margin, and a 
somewhat lesser angle with the straight tergo-lateral margin. There is no tergo-lateral slip formed by 
upturned lines of growth. Carina much bowed inwards, massive, with the basal margin bluntly pointed. 

Scania (Balsberg, Kopinge, Ffo., &c.) Mus. Univers. Copenhagen. 

Professor Steenstrup has described, under the name of Pollicipes Nilssonii, a large carina, and appa- 
rently a sub-carina and rostrum, and he remarks that these perhaps belong to the same species with the 
terga, named by him P. undulatus. M. Angelin, however, believes that the latter belong to the species 
already described as Scalpellum solidttlum. With the specimens of the present species, M. Angelin has 
lately found three small scuta, which he believes belonged to it. These scuta are so extremely worn, 
that I should not have ventured to have named them, had it not been advisable to give figures of the 
remarkable carina already named as P. Nilssonii. Should it hereafter be proved that the following scuta 
belong to some other carina, then a new name will have to be attached to them. 

Scuta (fig. 1 1, a) flat, thick, triangular, not much acuminated ; basal margin forming almost a rectangle 
with the occludent margin ; tergo-lateral margin (in present condition) straight, forming a rather less 
angle with the basal than does the occludent margin. There is no trace of a slip or portion of valve along 
the tergo-lateral side, formed by upturned lines of growth. Internally, the pit for the adductor muscle is 
deep ; the central portion of the apex above the pit is prominent ; apparently there was no internal furrow. 
Length of longest specimen only - 4 of an inch. 

Carina (fig. 11, b, c) strong, with the upper part unusually massive; though in a worn condition, 
there are distinct traces of its having been longitudinally and slightly ribbed. Strongly carinated, the two 
arched sides meeting each other at about a rectangle ; much bowed inwards, and widening much from 
the apex to the base ; upper portion, about one fifth of the entire length of the carina, seems (for the 
worn condition prevents certainty) to have projected freely; beneath the upper freely projecting portion, 
the inner margins are nearly straight ; the depth of the shell, measured from the central crest to the inner 
margin, is, in the lower half, remarkably great, and consequently the valve in the same part is internally 
concave to a remarkable depth ; the upper freely projecting portion is only slightly concave, and is singu- 
larly massive, from having been filled up with solid shelly layers. The basal margin is bluntly pointed, 
the edges meeting each other at about a right angle ; in the lower part of the valve the lines of growth 
are of course parallel to the basal edges, but higher up they meet at a more open angle, and consequently 
the carina of a young individual must have had its basal margin less projecting. When the sides of the 
carina are examined carefully, a portion, about one fourth of its entire depth, can be observed to lie a very 
little more inwardly inflected than the more central part, so as not to form quite a continuous surface with 



POLLICIPES. 53 

lower carinal margins compared with the upper carinal and occludent margins of the 
terga, and lastly in the basal margin of the carina being truncated ; it differs from P.planu- 
latus of the Oxford Clay, and therefore its other nearest relative in age, by the basal angle 
of the terga being rounded, instead of square as in that species. 

the two broad arched roof-sides ; and in these slips the lines of growth run almost parallel to the inner 
margin of the valve : in this respect the valve approaches in character to that of Scalpellum. The heels or 
baso-lateral angles apparently projected slightly, as I infer from a slight downward curvature in the lines 
of growth, along a line corresponding with the heel, and separating the roof-part from the inflected walls 
of the carina. 

Sub-carina (fig. 11, d).\ in Professor Steenstrup's collection there are several worn valves which appear 
to have been sub-carinse ; in shape approximately semi-conical ; the basal margin being almost semi-ovaL 
with the two comers a little inflected ; hence the valve is deeply concave to an unprecedented degree, and 
this is quite conformable with the singular sectional outline of the carina (c). About one fourth part of the 
length of the valve must have projected freely ; the outer surface is longitudinally ribbed, and the lines of 
growth remarkably undulatory. 

Rostrum (fig. 11, e) : this valve which I believe to be the rostrum resembles the sub-carina, but is more 
open, less high, and with a larger proportion, namely half, of its entire height freely projecting ; the semi-oval 
basal margin is slightly sinuous, the projecting points corresponding with the external longitudinal ribs. 

Length of carina, 1*5 ; of the largest of the sub-carinse, '6 ; of the largest rostrum, *45 of an inch. 

4. Pollicipes Hausmanni. Tab. Ill, fig. 3. 

Pollicipes Hausmanni. C. L. Koch and Dunker. Norddeutsch. Oolithgebilder, p. 52, Tab. vi, fig. 6. 

— — F.A.Roemer. Versteinerung. Norddeutsch. Oolithengebirges, p. 211, Tab. iv, fig. 2. 

P. scutis subtriangulis, angulo baso-laterali valde rotundato ; apice producto ; margine basali cum 
margine occludente angulum pane rectum formante ; internd apicis superjicie prominente, margineque 
tergali sulcato. 

Scuta, subtriangular, with the baso-lateral corner much rounded, and with the apex produced ; basal 
margin forming nearly a right angle with the occludent margin ; apex with its internal surface prominent, 
and with the tergal edge furrowed. 

Hilsthon, des Elligser Brinkes. (Lower Greensand, Germany.) 

Messrs. Koch and Dunker have given a full and detailed account of this species, together with truly 
excellent figures, and I have nothing to add to their remarks, but will re-describe, for the sake of uniformity, 
the valves of this species, which, through the kindness of Professor Steenstrup and Professor Dunker, 
I have examined. The valves are slightly worn. The figures given in tab. Ill are not, I think, so good as 
most of the others. 

Scuta (Tab. Ill, fig. 3, b, c) moderately convex, sub-triangular ; apex much acuminated, slightly 
curved towards the terga ; surface smooth, faintly marked with zones of growth, and, especially near the 
apex, with faint lines and furrows radiating from it. There is no distinct ridge proceeding from the 
apex to the baso-lateral angle, which is so much rounded that the basal margin blends into the tergo- 
lateral ; it must, however, be remarked, that the specimens are worn. The occludent margin' stands at 
right angles to the basal ; and the lower part of the tergo-lateral margin forms rather above a right angle 
with it. Internally (c), there is a deep pit for the adductor scutorum, and in the upper part, close to the tergal 
margin, a deep furrow ; the central portion is prominent ; the occludent margin keeps nearly of the same 
thickness up to the apex of the valve. 

Terga (Tab. Ill, fig. 3, d), nearly flat, sub-rhomboidal, or rather pointed oval, with the scutal half 



54 FOSSIL CIRRIPEDIA. 

5. Pollicipes politus. Tab. Ill, fig. 4. 

P. scutis fere rhombicis, lavissimis ; margine basali cum margine occludente angulum 
recto majorem formante ; margine occludente projecturd parietali? lineari, minutd instructo ; 
interna apicis superficie concavd. 

Scuta, almost rhoniboidal, excessively smooth, basal margin forming above a right angle 
with the occludent margin, which latter is exteriorly furnished with a linear, minute, wall- 
sided ledge ; apex with its internal surface concave. 

Mus. Bowerbank. Locality and formation unknown ; from the state of another specimen fastened on 
the same board, I think probably from the Gault ; the colour of the substance in the cracks of the valve 
countenances this same opinion. 

I have been unwilling to fix a specific name to a single, much broken scutum ; but as 
even in its present state it can be clearly seen to be distinct, and as this is the typical valve 
in this genus, I have felt myself compelled to do so. 

Scutum sub-rhomboidal, approaching to oval in outline : rather thin ; surface excessively 
smooth ; slightly convex, but with a narrow portion along the occludent margin, somewhat 
inflected : exteriorly close to this same margin, or rather almost forming it, (b) there is an 
extremely narrow, sharp, wall-sided, projecting ledge. The occludent margin is slightly 
arched, and forms, with the basal margin, an angle considerably above a right angle, so 
that the whole baso-lateral corner of the valve is much produced : the lower part of the 
tergo-lateral margin is at right angles to the basal margin. Baso-lateral angle smoothly 
rounded, with no trace of a ridge running from it to the apex, though this is the 
line of chief flexure of the valve. Internally, the valve has been much injured ; the de- 

protuberant ; surface smootb, but near the pointed, slightly curled apex, it is marked by fine radiating 
lines ; carinal margin regularly curved from the apex to the basal angle, which latter is not very sharp. 
A curved ridge (formed by the surface of the shell being lower on the carinal than on the other side) 
connects the upper and basal apices, running almost parallel to the carinal margin, and at about one-fourth 
of the entire width of the valve from the latter margin. Occludent margin shorter than the scutal ; 
rounded, protuberant, with a depression parallel to it; the scutal margin, corresponding with this depression, 
being slightly hollowed out ; a small portion of the apex projects freely. Internally, and nearer to the 
occludent than to the carinal margin, there are three or four short parallel longitudinal ridges or crests, as 
described in Scalpelhon arcuatum. 

Carina (Tab. Ill, fig. 3, a) moderately bowed inwards, widening gradually from the apex to the 
basal margin, which is rounded and protuberant, and with a trace of an angular bend in the middle ; 
exteriorly the surface presents just a trace of being sub-carinated ; roof convex ; the upper part of the 
valve projects freely. 

Rostrum : Koch and Dunker figure valves, which, from their general appearance, breadth, and appa- 
rently large proportional upper, freely-projecting portion, I have little doubt have been rightly considered 
by them as rostra ; they are, however, longitudinally plicated or striated to a greater extent than the 
other valves. 

1 Parietali, i. e. lateribus utrinque perpendicularibus. 



POLLICIPES, 55 

pression for the adductor scutorurn does not appear to have been deep : the concavity of 
the valve extended to the apex, with the upper part not filled up solid ; a considerable 
portion of the upper tergal margin is marked by lines of growth, and must have over- 
lapped the tergum, but there is no trace in it of a recipient furrow. 

Affinities. This species seems quite distinct from all others ; and I can hardly say to 
which species it is most related ; in some few respects it comes nearest to P. acuminatus} 

1 6. Pollicipes ELONGATUS. Tab. Ill, fig. 5. 

Pollicipbs elongatus. Steenstrup. Kroyer's Tidsskrift, b. i, p. 361. 

— l^vis (Sowerby). lb. lb. b. ii, p. 409, pi. 5, figs. 9, 10. 

P . scutis pcene quadratis ; margine occludente et parte inferiori marginis tergo-lateralis rectangule ad 
marginem basalem positis ; apice obtuso. 

Scuta almost square ; occludent margin and the lower part of the tergo-lateral margin at right angles 
to the basal margin ; apex blunt. 

White Chalk, Denmark. Mus. Univers. Copenhagen. 

General Remarks. Professor Steenstrup at first described this species as distinct, but subsequently con- 
sidered it the same with P. Icevis of Sowerby ; this is not the case, and therefore I have retained the name 
first given, though very inappropriate to the more important valve. According to the practice here followed, 
the species is founded on the scutum, of which a mere fragment and impression exists, but it is sufficient 
to show that it is distinct. In the same chalk with this scutum, there are two terga which are different 
from any other seen by me, and which Professor Steenstrup has described as belonging to this species, a 
view which I have followed with some hesitation. 

Scutum (fig. 5, c, b), nearly smooth, but with the growth-lines plain ; thin, very slightly convex ; oblong, 
almost rectangular, but with the upper lateral corner rounded off. Occludent and lower part of tergo-lateral 
margin both straight, and at right angles to the straight basal margin : upper part of tergo-lateral margin 
much arched and protuberant, which, together with the blunt apex given to the valve, it is almost rectangular, 
oblong outline. A trace of a ridge runs from the baso-lateral angle to the blunt apex. The figure 
given is a restoration ; being guided by the outline impressed on the chalk, and the lines of growth as seen 
on the small baso-lateral preserved portion. 

Tergum (fig. 5, a), convex, thick, elongated, sub-rhomboidal, or rather triangular, for the upper and lower 
carinal margins blend into each other with quite a uniform and gentle sweep : upper part of the carinal and 
occludent margins meet at an angle of about 45° ; from the apex to the sharp basal angle, an angular, very 
slightly curved, conspicuous ridge runs at about one third of the entire width of valve from the carinal 
margin : the surface of the valve slopes rather steeply away on both sides from this ridge. Occludent and 
scutal margins about equal in length ; parallel to the former, a rather wide space of the valve is slightly 
depressed, with a corresponding portion of the scutal margin slightly indented ; the occludent margin 
itself is not rounded and protuberant, as if it had been received in a furrow in the scuta. A considerable 
portion of the upper part of the valve projected freely. This valve more resembles the homologous one in 
the Italian tertiary, P. carinatus, than that of any other species : as before stated, I assign it to this species 
on the excellent authority of Professor Steenstrup ; I may, however, remark, that it appears much thicker 
and stronger than the scutum. The figure of the tergum (a) is not very good, and has been drawn on too 
small a scale. 



56 FOSSIL CIRRIPEDIA. 

7. Pollicipes acuminatus. Tab. Ill, fig. 6. 

P. scutis elongatis, triangularis ; margine basali cum margine occludente angulum recto 
longe minorem formante ; interna apicis superjicie concavd. 

Scuta elongated, triangular ; basal margin forming much less than a right angle with 
the occludent margin ; apex, with its internal surface concave. 

White Chalk. Mus. Flower, (believed to have come from the Lower Chalk of Stoke Ferry, Norfolk). 

My materials consist only of a single left-hand scutum, and that with the whole of the 
basal margin broken off; nevertheless, there can be no question that it is quite distinct 
from all the species hitherto described. 

Scutum : shell rather thin, surface extremely smooth ; triangular, much elongated, with 
the upper part a little bent over towards the terga : slightly convex, but with the whole 
middle part of the valve remarkably flat; the convexity being caused by the inflec- 
tion, in a slight degree, of the occludent margin, but chiefly of the tergo-lateral portion ; 
hence, a smooth ridge of chief curvature runs from the apex to the baso-lateral angle. 
Occludent margin arched, forming less than a rectangle with the straight basal margin, 
with which the very slightly concave tergo-lateral margin forms an angle rather above a right 
angle : the tergo-lateral portion of the valve, formed by the upturned zones of growth, mode- 
rately wide, being in the upper part about one third of the entire width of the valve : 
rostral angle rounded. Internally (b) the valve is singular ; the depression for the adductor 
scutoruni muscle is extremely faint, and is situated unusually low down in the valve, though 
the exact relative position must at present remain unknown, as the basal margin has been 
broken off. But the most unusual character, at least in the cretaceous species of Pollicipes, 
is, that the concavity of the valve runs up to the apex, and must have been lined up to that 
point with corium : the flat internal occludent edge, marked by lines of growth, widens 
very little in the uppermost part. 

8. Pollicipes angelini. Tab. Ill, fig. 7. 

P. scutis elongatis, triangulis, margine basali pjrope angulum rostralem in prominentiam 
oblique rotundatam producto : interna apicis superjicie prominente, margine occludente 
sulcato. 

Scuta elongated, triangular, with the basal margin near to the rostral angle, produced 
into an obliquely rounded point ; apex with its internal surface prominent, and with the 
occludent edge furrowed. 

Upper Chalk, Norwich. Mus. Fitch. Kjuge, Scania. Mus. UniverR. Copenhagen. 

My materials consist of four scuta which, as usual, I take as typical, a pair of terga, 
and one or two carinse which, for reasons to be given, perhaps belonged to this species ; all 



POLLICIPES. 57 

in Mr. Fitch's Collection : also a single scutum sent me by Professor Steen strap, and found 
by M. Angelin, after whose name I have called this well-marked and peculiar species. 

Scuta (fig. 7, a, b) ; triangular, much elongated, considerably convex, apex extremely 
acuminated ; basal margin at nearly right angles to the straight occludent margin, but near 
to the rostral angle, it is produced in a remarkable manner into a rounded, obliquely truncated 
broad point. 1 The tergo-lateral portion of the valve, formed by the upturned lines of growth, 
is not much developed : the tergo-lateral margin, as seen externally, is obscurely divided 
into two lines, of which the upper, or tergal portion, has its edge reflexed ; this same whole 
margin, however, seen internally, appears nearly straight, and this is essentially the case ; 
the projecting angle being connected with the thickening of the valve during growth. The 
exterior surface is smooth, with some faint longitudinal striae : a single ridge, or rather, 
line of flexure, runs from the apex to the baso-lateral angle. Seen internally, the upper- 
most part of the valve is found to be unusually thick and solid, with the pit for the ad- 
ductor scutorum muscle well developed, and placed rather low down. The internal occlu- 
dent edge (3), marked with lines of growth, becomes close above the adductor depression 
suddenly very wide, and forms a deep furrow, which I at first thought was formed to re- 
ceive the occludent angle of the terga ; but upon consideration, I feel pretty sure that this 
cannot have been the case, and I believe the furrow to be of no functional importance, but 
to result from the sharp apex of the still corium-covered portion of the valve having been 
greatly thickened : this same upper portion has, in most specimens, in its middle, a slight 
linear furrow. On the tergal margin of the internal surface there is a small portion, marked 
with lines of growth, which is obliquely truncated, owing to the valve having become very 
thick; and this must have overlapped the tergum. From these peculiarities in the internal 
surface of the apex of the scuta, it may be inferred, that the terga, owing to the probable 
close contact of the two valves, would present peculiarities of a corresponding nature. 
The largest British specimen is "8 ; and the Scanian specimen is "95 of an inch, in length. 

Terga (fig. 7, c, d) ; in Mr. Fitch's collection there are, from the same formation in 
which the above scuta were found, two terga, remarkable from a very wide square-edged 
depression, running parallel to the occludent margin, which is itself rounded and pro- 
tuberant : these valves probably belonged to the P. Angelini, and anyhow may be con- 
veniently here described. Valve rhomboidal, not very flat, smooth, with a conspicuous 
ridge, wall-sided on its carinal aspect, running almost down the middle of the valve from 
the apex to the basal angle, which latter is not very sharp. The upper carinal and 
occludent margins meet each other at slightly less than a right angle ; occludent margin 
a little longer than the scutal margin, with its edge thickened, rounded, and protuberant 
to an unusual degree ; alongside the occludent margin an unusually broad and deep, 
square-sided depression runs, equalling in width about one third of the scutal margin ; a 

1 Amongst the Scanian fossils from Kopinge, there is a scutum with the whole upper part broken off, 
but which I think belonged to this species ; if so, it differs from all the others in the rostral portion of the 
basal margin projecting yery little. 

h 



58 FOSSIL CIRRIPEDIA. 

transverse section across the middle of the valve is given in fig. 7, d, exhibiting its peculiar 
outline. 

Carina. In Mr. Fitch's collection there are two carinas which probably belonged to 
this species, at least they do not belong to the only two other species of Pollicipes, (viz. 
P.fallax and striatus,) found by this gentleman in the chalk near Norwich. These carinas 
are identical, as I know from examination, with that (also from Norwich) figured by 
Mr. Sowerby, (Min. Conch., PI. 606, fig. 7,) as the anterior valve of his Pollicipes 
sulcatus, Scalpellum maximum, var. sulcatum of this work. These two carinas differ a 
little from each other in the basal margin, being either rectangularly pointed or rather 
blunter and more arched ; their surfaces are smooth, but in one specimen there are 
some obscure longitudinal strias ; in outline they are tapering, triangular, almost straight, 
transversely flatly arched, subcarinated, edge inflected, with the lines of growth bowed 
downwards, and consequently with the basal lateral angles or heels a little prominent. 
Finally, these carinas cannot be strictly distinguished from the same valve in P. glaber 
and unguis, but as I believe neither of these species occur in the upper chalk near Norwich, 
they can hardly belong to them. 

Affinities. The scuta in this species differ from all others in the projection of the basal 
margin, close to the rostral angle ; a somewhat similar projection, I may remark, sometimes 
occurs in the scuta of the recent Lepas fascicularis, A. vitrea of Lamarck. Internally, the 
valve more resembles that of the second and third varieties of Scalpellum maximum than 
any other species. The terga can be distinguished from those of P. glaber only by 
the depression along the occludent margin being deeper, wider, and square-sided. 



9. Pollicipes reflexus. Tab. Ill, fig. 8. 

Pollicipes reflexus. /. Sowerby. Min. Conch., pi. 606, fig. 8. 

P. scutis tenuibus, subovalibus ; margine basali cum margine occludente angulum reck 
longe majorem formante ; costd obsoletd rotundatd ab apice ad angulum baso -later alem decur- 
rente, valvam in duas fere cequales partes separante. Carina lineari, transverse abrupte 
arcuatd ; margine basali multurn jjroducto, apice truncate-. Lateribus superioribus sub- 
pentagonis. 

Scuta ; thin, suboval, with the basal margin forming an angle much larger than a rect- 
angle with the occludent margin ; a faint rounded ridge runs from the apex to the baso- 
lateral angle, dividing the valve into two nearly equal halves. Carina linear, in a trans- 
verse line steeply arched ; basal margin much produced, with its apex truncated. Upper 
Latera subpentagonal. 

Upper Marine Eocene formation ; Colville Bay, Isle of Wight. Mus. J. Sowerby, F. Edwards. 
I owe to the kindness of Mr. Sowerby an examination of some of the original speci- 



POLLICIPES. 59 

mens, consisting of a scutum, terga, carinse, and upper latera ; and to Mr. F. E. Edwards 
the loan of several very perfect specimens. 

Valves ; smooth, rather thin. Scutum (fig. 8, e) elongated, very slightly convex, many- 
sided, almost oval, with its upper point produced, but apparently apt to be broken off. 
Occludent margin slightly arched ; basal margin consists of two sides, of which the rostral 
side is extremely oblique to the other, the two sides together forming a very large angle 
with the occludent margin ; the tergo-lateral margin also consists of two sides, meeting 
each other at a very open angle ; the lower half forms above a rectangle with the basal 
margin ; possibly this lower half corresponds with the widely-truncated baso-lateral angle 
in P. dorsatus, and with the smaller similar angle in the tertiary P. carinatus. A faint 
rounded ridge runs from the apex to the baso-lateral angle, and thus divides the valve into 
two nearly equal halves. Internally there is a deep pit for the adductor muscle ; only a 
narrow internal edge along both sides of the apex is marked by lines of growth. 

Terga (fig. 8, d) ; elongated, rhomboidal, flat and thin ; carinal margin continuously 
curved; apex much produced; the occludent and upper carinal margins meet each 
other at much less than a right angle ; occludent margin slightly longer than the scutal 
margin, which latter in the upper part is very slightly hollowed out ; a faint, rather wide 
ridge, runs down the centre of the valve to the rectangularly-pointed basal angle ; a small 
portion of the upper part of the terga projected freely. 

Carina (fig. 8, a, b, c) ; narrow, with the lower part somewhat spear-shaped ; almost 
straight, transversely convex, with the central part forming a broad rounded crest ; lateral 
margin a little inflected; basal margin extraordinarily produced, with its apex square; 
internally, the corium- covered surface is produced upwards into the sharpest apex; a 
small portion only of the uppermost part of the valve projected freely. 

Upper Latera (fig. 8, /) ; flat, almost pentagonal, with the two upper margins consider- 
ably longer than the three lower ones ; of these three, the carinal margin is the shortest, and 
the other two equal, with the rostral one considerably arched or convex ; these three basal 
margins must have been in contact with the Latera of the lower whorl ; the apex did not 
project freely. I have not seen any latera of the lower whorl, but Mr. Sowerby gives 
figures of some minute valves, which no doubt were such. Several of Mr. Edwards's speci- 
mens are tinged a pale-reddish purple. 

Dimensions. The valves described are all rather small ; the largest, a tergum, is a little 
more than a quarter of an inch in length. 

Affinities. This species is more nearly related to the section of the genus containing the 
recent P. cornucopia, elegans, and pjolymerus, than to the other sections. Its affinity is 
closest to P. cornucopia, though in the form of the basal margin of the scuta there is more 
resemblance to P. polymerus. The scutum presents some points of resemblance to the 
cretaceous P. gracilis and dorsatus. 



60 FOSSIL CIRRIPEDIA. 

[i?.] Scuta, costd, nonnumquam subobsoZetd, ab apice ad centrum marginis basilis 
decurrente} 

1 10. Pollicipes carinatus. Tab. Ill, fig. 9. 

Pollicipes carinatus. PhUUppi. Enum. Mollusc. Siciliae, 1836, Tab. xii, figs. 26, 28. 

P. scutis crassiusculis ad formam trianguli cequianguli accedentibus ; margine occludente externe costd 
humidi firmato ; costd fir md ab apice ad centrum marginis basulis decurrente ; margine basalt recto; angulo 
baso-laterali truncato, brevi ; segmentum t ergo-later ale ex lineis incrementi reflexis formatum, deest. Carind 
externe valde carinatd cum sulco laterali ad utrumque latus. 

Scuta moderately thick, in shape nearly an equilateral triangle ; occludent margin exteriorly strengthened 
by a slight ridge ; a strong ridge runs from the apex to the middle of the basal margin ; basal margin 
straight ; baso-lateral angle truncated, short : the tergo-lateral portion, formed by upturned lines of growth, 
is absent. Carina, exteriorly, strongly carinated, with a lateral furrow on each side. 

Tertiary ; Messina, Sicily. 

I owe to the great kindness of Dr. Phillippi an examination of an authentic series of specimens. Valves 
rather thick, with the lines of growth plain, and with a few fine striae radiating from their apices : these 
striae are sometimes so conspicuous, that I have doubted whether the species ought not to have been 
placed in the next section. 

Scuta (fig. 9, d, e) ; triangular, with the apex slightly bowed over towards the terga ; moderately convex ; 
occludent margin slightly arcbed, about equal in length to the slightly hollowed-out tergo-lateral margin ; 
basal margin nearly straight, though formed by two lines meeting each other, sometimes with the 
rostral half not descending so low as the other half; this margin forms equal angles with the other two 
margins. The baso-lateral angle is obliquely truncated : there is no tergo-lateral slip formed by upturned 
zones of growth. The occludent margin is exteriorly strengthened in a manner only just perceptible 
by a flattened rim. A strong, conspicuous, and prominent ridge runs, in a slightly curved course, from 
the apex to a point in the basal margin, rather nearer to the rostral than to the baso-lateral angle ; 
this point just perceptibly projects beyond the rest of the basal margin : the ridge is either moderately 
sharp, or broad and flat-topped ; it marks the line of chief curvature of the valve. Internally (e), the pit for 
the adductor muscle is not very strongly developed, and to a different degree in different specimens: the 
internal surface of the centre of the apex is prominent ; on its tergal side there is no furrow, but a rect- 
angular indentation formed by a remarkably wide, flat, smooth ledge, which runs down, narrowing, to the 
baso-lateral angle ; hence the scuta along the whole of this side, especially in the upper part, must have widely 
overlapped the terga, in a manner and to a degree I have not seen equalled in any other Pollicipes; but the 
two valves cannot be said to have been articulated together. The internal occludent edge widens a little in 
the upper part, and is here divided by an oblique line, with the lines of growth apparently discontinuous on 
opposite sides of it, into two portions, of which the inner portion is slightly more prominent than the outer. 
I have already alluded to the fact, that in some specimens the scuta are strongly ribbed longitudinally, 
in some very faintly striated, and in others smooth. 

Terga (fig. 9, a, b) ; sub-rhomboidal, elongated, exteriorly convex, internally very slightly concave : upper 
part very thick and solid, with a considerable portion freely projecting, and internally marked by lines of 
growth : along the middle of the portion thus marked, there is a slight longitudinal depression, which is worth 
remarking, inasmuch as (judging from a conspicuous and analogous character in certain recent species of 
Lithotrya) it was probably caused by the internal central crest of the upper part of the carina. Occludent 
margin slightly convex, very little shorter than the scutal margin ; upper and lower carinal margins nearly 
equal in length ; they meet each other at a very open angle : upper carinal and occludent margins meet each 
other at an angle of only a little above 50°. A prominent, strongly marked ridge, (with both sides of the 



POLLICIPES. Gl 



11. POLLICIPES GLABER. Tab. Ill, fig. 10. 

Pollicipes glaber. Roemer. Norddeutsck. Kreidegebirg. Tab. xvi, fig. 11. 
Xiphidium maximum. J. Sowerby. Dixon's Geology of Suffolk, Tab. xxviii, figs. 6 — 8. 

P. seutis subtennibus, latiusculis, adformam trianguli cequianguli accedentibus ; margine 
basali non prorsus recto; tergo-laterali segmento e zonis incrementi reflexis formato, 
ubi latissimo, reliqua valval dimidium aquante : margine tergali apicis intus sulcata. 
Carina margine basali obtuse acuminate-. Lateribus superioribus triangulis, tertiam partem 
longitudinis tergorum cequantibus. Lateribus anticis inferioribus singulis costd prope termi- 
num marginis basal is decur rente. 

valve sloping from it,) runs from the apex to the sharp basal angle : this ridge is very slightly curved in 
two directions, like the letter S, it runs at about one third of the entire width of the valve from the 
carinal margin. 

Carina (fig. 9, e,f) strong and solid, with lines of growth conspicuous on its surface ; very slightly arched 
inwards ; triangular, moderately tapering ; transversely moderately convex, very plainly carinated, with a 
slightly projecting rib ; on each side, at a little distance from the lateral edges, there is a distinct and linear 
furrow, and these edges themselves are, partly in consequence, rounded and slightly protuberant : basal 
margin square, and not at all protuberant. Internally, the upper part, for about one third of the total length 
of the valve, must have projected freely, and has been filled up solid with a trace (/) of a central crest : 
the internal lateral edges are slightly scalloped out along the whole length of the valve. 

Rostrum (fig. g, h, i) ; broad, triangular : apex curled inwards to a most remarkable extent, so as even to 
point a little downwards ; basal margin just perceptibly protuberant, with a square projection formed by the 
end of a wall-sided, broad, flat-topped ridge, running down from the apex ; Phillippi, however, states, that 
the form of the basal margin varies. Internally, a full upper half of the valve projected freely ; the internal 
upper surface is smooth and concave, with just a trace of a fine central crest: the inwardly curled apex 
converts the upper part into a hood : the central basal projection is channelled, the channel running a little 
way up the valve, and being gradually lost : this channel, no doubt, allowed a filament of the corium to pass 
to the sub-rostrum. 

Peduncle. Dr. Phillippi has sent for my inspection, a rare and interesting specimen of a peduncle, 
with the scales preserved, no doubt, belonging to this species. The scales, as usual, decrease downwards in 
size ; they are rather broad ; each has its upper end rounded ; is marked transversely by lines of growth, 
and has a slight external, central, longitudinal crest : this crest is wedge-formed, being widest at the 
apex. I have not seen this latter character in the peduncular scales of any other Pollicipes ; there are, how- 
ever, traces of it in the small lower Latera in P. mitella; it is apparently caused by the lateral overlapping 
of the closely-packed scales, and chiefly when the specimen was young. 

Affinities. The general form of the carina, with its lateral furrows and rounded protuberant margins, — its 
carinated central line ; — the shortness, and great inward curvature of the rostrum, with its strong, central 
ridge, terminating in a channelled projection on its basal margin — show a clear affinity between this species 
and the recent P. mitella of the Eastern tropical seas. There are some points of resemblance in the scuta and 
terga to the same recent species ; but in the scuta a closer affinity is shown to P. dorsatus ; and in the terga 
to those described under P. elongatus : in the manner, moreover, in which the upper part of the carina is 
filled up and furnished with a central crest, there is a relationship to P. validus. The affinity of P. carinatus 
to P. mitella is interesting ; because M'e may with some confidence infer from the relationship between 
P. carinatus, dorsatus, validus, and rigidus, that, in these several species, the valves, which hitherto have 
been found only separated, were united together to form the capitulum in a somewhat similar manner as in 
the well-known recent P. mitella. 



62 FOSSIL CIRRIPEDIA. 

Scuta moderately thin, rather broad, approaching to an equilateral triangle ; basal 
margin not quite straight ; the tergo-lateral portion, formed by the upturned zones of 
growth, where widest, half as wide as the rest of valve ; apex internally furrowed on the 
tergal side. Carina with the basal margin bluntly pointed. Upper latera triangular, one 
third the length of the terga. Anterior lower latera, each with a ridge running to near 
one end of their basal margins. 

Lower Chalk, Stoke Ferry, Norfolk. Lower Chalk, Hanover, according to Roemer. Planer (Chalk- 
marl), Sarstedt, near Hildesheim, Mas. Roemer. Upper Chalk, Northfleet and Gravesend, Kent. Chalk 
Detritus, Charing, Kent. Maastricht Formation, Scania? Mus. Flower, Wetherell, Harris, Univers. Copen- 
hagen, Geolog. Soc, and Bowerbank. 

General remarks. My materials consist of several scuta, in Mr. Flower's collection from 
Stoke Ferry, together with some carina?, and a rostrum of apparently the same species ; of 
a single scutum from Northfleet, and of others from Charing, both in Kent ; of some scuta, 
terga, and carinse, sent me by the great kindness of Roemer, from Sarstedt, near Hildesheim, 
and therefore authentic specimens by him named ; and lastly, of a valuable specimen from 
Gravesend, in Kent, in Mr. Bowerbank's collection (fig. 10, a), in which a carina, pair of 
terga, an upper latus, and two lower latera, were embedded in nearly their proper positions, 
together with a fragment of a scutum, which latter is sufficiently perfect to leave no doubt on 
my mind regarding its identity with the valves from Stoke Ferry, Northfleet, and Sarstedt. 
This appears to have been one of the commonest species of Pollicipes during the cretaceous 
period. Though found in the upper chalk of Northfleet and Gravesend, it is singular that 
not a single valve of this species has been collected by Mr. Fitch in the upper chalk of 
Norwich. Amongst the Scanian fossils, collected by M. Angelin at Kopinge, from a still 
higher stage of the chalk, and forwarded to me by Professor Steenstrup, there is an upper 
latus and tergum most closely allied, probably even identical with the present species. 
We have seen that it extends down even to the planer or chalk marl. 

Scuta (fig. 10, b, c, d). These valves are moderately thick and convex, so that in their 
upper halves they are almost semi-conical ; the basal margin is not quite straight, it forms a 
rather larger angle with the lower part of the tergo-lateral margin, than with the occludent 
margin, both angles being less than right angles. From the apex two faint ridges run, 
one to the baso-lateral angle, and the second to a point in the basal margin, a little nearer 
to the rostral than to the baso-lateral angle. The valve is bent, so as to be convex, chiefly 
along these two ridges. The tergo-lateral portion formed by the upturned lines of growth 
is wide and protuberant. Outer surface of valve smooth, with the faintest striae radiating 
from the apex. Viewed internally [d), a conspicuous furrow runs from the pit for the 
adductor scutorum muscle up to the apex ; the internal occludent edge keeps the same 
width up to the apex. The Sarstedt specimen is the largest scutum which I have seen, 
and that is - 6 of an inch in length. 

One out of Mr. Flower's five specimens (believed all to have come from the lower 
chalk of Stoke Ferry) and another in Mr. Wetherell's collection from the upper chalk of 



POLLICIPES. 63 

Northfleet, present some slight differences, in the valve being flatter, in the tergo-lateral 
portion being more developed, and in the basal margin being straighter ; but these differ- 
ences are so very slight that it would be exceedingly rash to consider them as specific. 

Terga (fig. 10, e, a) ; rhomboidal, flat, with a straight slight ridge running, at about one 
third of the entire width of the valve from the carinal angle, to the sharp basal angle ; this 
ridge is steep on its carinal side, on which side the whole surface of the valve is somewhat 
depressed. The upper carinal margin meets the occludent margin at somewhat less than a 
right angle: the occludent margin slightly exceeds in length the scatal margin; the valve 
is widest nearly in its middle. A narrow rim along the occludent margin is slightly pro- 
tuberant, within which there is a slight parallel depression. The scutal margin is not quite 
straight ; the lower half (which probably was in contact with the upper latus) projecting a 
little. 

Carina (fig. 10,/,^); triangular, tapering, moderately strong, with a smooth surface; 
almost straight or slightly curved inwards ; transversely, flatly arched, sub-carinated. 
Lateral margins narrow, inflected, with the lines of growth first curved downwards (g), 
and then on the edge itself abruptly upturned ; basal margin bluntly pointed, apparently 
in a variable degree. The two baso-lateral angles are slightly prominent, in conformity 
with the above-stated direction of the lines of growth along the margin. I may remind 
the reader that this valve, the terga, and the latera presently to be described, were all 
embedded together in their proper positions. 

Mostrum (fig. 10, h, i) : in Mr. Flower's collection, with the other loose valves from 
Stoke Ferry, there is one which I can hardly doubt is a rostrum : it is nearly an equilateral 
triangle ; externally convex, sub-carinated, basal margin not protuberant. The upper part, 
for about one fourth of the entire length of the valve, must have projected freely ; this 
part is remarkable, from having been filled up internally with a central crest, like the carina 
of P. validus and carinatus. 

Upper latus (fig. 10, h, a) ; flat, almost an equilateral triangle, but with the two upper 
sides not exactly equal, and both a very little longer than the basal margin ; umbo 
of growth at the uppermost point ; length, one third of the terga ; surface smooth, with no 
trace of a central longitudinal ridge ; internally the edges are bevelled, and a very small 
portion of the apex must have projected freely. 

Lower latera (fig. 10, /, a). There are two of these (one much broken) nearly resembling 
each other : from analogy with the latera of P. unguis, these are probably from the rostral 
end of the capitulum ; they are slightly convex, and approximately form a transversely 
elongated triangle, with one corner cut off ; the two upper sides are slightly unequal in 
length : umbo of growth at the apex, from this point a narrow well-defined ridge runs 
obliquely across the valve, to the most protuberant point of the basal margin, which is 
situated about one fourth of the entire width of valve from one end. These lower latera 
are wider than, but not so high as the upper latus : seen internally the edges are bevelled, 
and a very small portion of the apex must have projected freely. 

As it is so very rare to find the valves of a Pollicipes united, it will be advisable to give 



64 FOSSIL CIRRIPEDIA. 

the measurements of the Gravesend specimen : the scutum is too much broken to be 
measured, and the breadth only of the basal margin of the carina can be given — it is "25 of 
an inch ; the terga are nearly '8 long and fully '4 broad. The upper latus is "27 in height, 
basal margin "23 in length. Lower latus *2 in height and "26 in breadth. I imagine 
that the broken scutum belonging to this individual, was fully two thirds of the size of the 
largest scutum which I have seen, namely, that from Sarstedt, *6 of an inch in length. 

JJinities. From the very close affinity of this species to the following one, P. unguis 
of the Gault, we may safely infer that it had not only a rostrum (of which a specimen found 
separately has been described), but also a sub-rostrum ; whether it had a sub-carina, as is 
probable, must remain doubtful. The two faint ridges running from the umbo of each 
scutum to its basal margin, probably mark the extent to which this valve was overlapped, 
as in the case of the recent P. mitella, by the rostrum and upper latera; and the ridge on 
the terga, steep towards the carina, probably shows the extent to which this latter valve 
reached. It is also probable, as we shall see under the head of P. unguis, that there was 
only one whorl of valves under the upper latera, and that this included the sub-rostrum : 
the latera in this lower whorl were probably not very numerous, but large, and of very 
unequal sizes ; their mutual overlapping no doubt caused the oblique ridges on their 
exterior surfaces. 



12. Pollicipes unguis. Tab. IV, fig. 1. 

Pollicipes unguis. J. Sowerby. Geolog. Transac. 2d series, vol. iv, t. xi, fig. 5*. 

— l.evis. lb. lb. lb. fig. 5 (sed 

non Tab. xvi, fig. 1, — alia species). 

P. scutis incognitis, verisimiliter ut in P. glabra. Margins basali carina obtuse 
acuminato, sub-rotundato. Lateribus superioribus[ elongdtis triangularibus, dimidium lon- 
gitudinis tergorum siiperantibus. Lateribus anticis inferioribus costd subcentrali instructis. 

Scuta unknown, probably as in P. glaber. Carina, with the basal margin, bluntly 
pointed, somewhat rounded. Upper latera elongated, triangular, exceeding half the 
length of the terga. Anterior lower latera, each with a ridge, subcentral. 

Gault ; Folkstone, (common.) Burham, Eastweare Bay, Maidstone. Mus. Practical Geolog., Bower- 
bank, J. Sowerby, Brit. Mus., &c. In the Museum of Geolog. Soc. there is a tergum of this species, 
marked Lower Greensand, Maidstone. 

General Bemarfo. This species comes very close to P. glaber, but can, I think, be 
safely distinguished from it, even with our present materials. A specimen most unusually 
perfect is in the Museum of Practical Geology ; it consists of a carina and pair of terga, 
much mutilated, a rostrum, sub-rostrum, a pair of upper latera, a pair of latera of the 
lower whorl from the carina! end of the capitulum, and two other latera of this same 



POLLICIPES. 65 

whorl from one side of the rostral end of the capitulum. All these valves are magnified 
twice in fig. 1, Tab. IV ; and as all, except (d), belonged to the same individual, we here 
have the rare advantage of learning their relative sizes. The largest of the lower latera has 
been mistaken for a scutum, and has been thus figured by Mr. Sowerby ; the mistake was 
a very natural one, to be rectified only by examining the under side of the specimen. 
Although the scutum is, unfortunately, at present unknown, there can be scarcely 
any doubt that it would closely resemble that of P. glaber, and therefore I have not hesi- 
tated, in this instance, to break through my rule of exclusively taking the scutum as typical 
in Pollicijpes : should, hereafter, a scutum be found in the Gault like that of P. glaber, it 
may, with considerable confidence, be named as belonging to this species. 

I have felt considerable doubts regarding the nomenclature of this species : P. unguis 
is founded on one of the latera of the lower whorl, and on what apparently is a rostrum ; 
these valves are at present, and will probably for very long remain insufficient for the 
foundation of a species. P. lavis is founded on a tergum and carina, and therefore on 
better grounds. The specimen immediately to be described, leads me to believe that all 
these valves belong to the same species ; and therefore both names were open to me. 
Mr. Sowerby, however, has given in the same volume of the ' Geological Transactions,' the 
name of P. leevis to a carina and tergum from Blackdown, which, I cannot doubt, is distinct : 
Professor Steenstrup has also described a new form under the same name of Icevis ; 
such being the case, it has appeared to me advisable to take the name of P. unguis. I 
must add, that there is some considerable variation in the terga from the Gault, which 
renders it just possible, but not probable, that there may be a second closely allied form. 
It is very singular, considering how very frequent terga are in all collections, that I should 
not have seen a single scutum which could, as I believe, have belonged to this form. 

Scutum unknown. Tergum (Tab. IV, fig. 1, b, c); this is a medium-sized specimen, 
•5 or "6 of an inch in length; it is rhomboidal, nearly flat, with a straight, slight ridge, 
running from the apex, at about one third of the entire width of the valve from the carinal 
angle, to the sharp basal angle : the ridge is steep on its carinal side, on which side the 
whole surface of the valve is somewhat depressed. The upper carinal margin meets the 
occludent margin at somewhat less than a right angle. Internally, a rather small portion 
of the apex is marked by lines of growth ; and close along the upper carinal margin there 
is a narrow furrow (b), with a ridge parallel and exterior to it ; both furrow and ridge are 
rather variable, and, no doubt, are produced by contact with the edge of the carina. The 
occludent margin very slightly exceeds in length the scutal margin ; the valve is widest 
nearly across the middle. A slight depression, with just perceptibly angular sides, runs 
parallel to the occludent margin. The scutal margin is nearly straight ; with the exception 
of this latter single character, scarcely any difference can be perceived between this valve 
and that of P. glaber. 

I have seen several large terga, (d, natural size,) nearly an inch in length, from the 
Gault, which at first appeared so different, that I thought them specifically distinct ; they 



66 FOSSIL CIRRIPEDIA. 

form the variation above alluded to. But, after careful comparison of a large series, I feel 
convinced that the difference is caused solely by age : the chief difference consists in the 
occludent margin being considerably longer than the scutal margin, and consequently 
in the widest part of the valve lying below the middle point : that portion of the valve, 
moreover, which lies on the carinal side of the sub-central ridge is, in proportion, narrower 
than in the common form. Internally, in these large specimens, a considerable portion of 
the apex is marked by lines of growth : it follows from this, that if only the internal 
corium-covered surfaces of the large and small terga be compared, there is far less 
difference of outline than if they be compared externally. I repeat that I have no doubt 
that these valves all belonged to the same species. 

Carina (Tab. 4, fig. la), closely like that of P. gldber, triangular, moderately tapering, 
very slightly bowed inwards ; smooth, sub-carinated, transversely slightly convex ; lateral 
margins very narrowly inflected, with the lines of growth curved downwards, and those on 
the edge itself upturned ; there is in this specimen a narrow, slight, linear channel along 
the line where the lines of growth are upturned ; a small portion of the apex must have 
projected freely ; basal margin (and consequently lines of growth) rounded, less angularly 
protuberant than in P. gldber, with the two baso-lateral angles slightly prominent. 

Rostrum (fig. le), triangular, rather more than half as wide and about half as long as 
the carina, therefore rather wider in proportion to its length ; more bowed inwards ; a very 
small portion of the apex, which is internally simply concave, projected freely ; basal margin 
curved, slightly protuberant : exterior surface smooth, sub-carinated, like the carina ; edges 
very thin. Sub-rostrum (/) about half as long, and two thirds as wide as the rostrum ; 
apex rounded ; basal margin not protuberant ; internally, slightly concave, with thin lateral 
margins, widely overlapping the latera of the lower whorl ; exterior surface not sub- 
carinated, that is, destitute of a central ridge. 

ZJ/jper Latera (jf), elongated, triangular, flat, exteriorly smooth, except from the lines 
of growth ; two upper sides almost exactly equal in length, and equalling once and a half 
of the length of the basal margin ; entire valve rather exceeding half the length of the 
terga. 

Lower Latera : these consist of two small valves (/, k), namely (judging from the 
position in which, overlapping each other, they were embedded), the first and second, or 
more probably the second and third right-hand rostral latera of the lower whorl ; and a 
pair {It, i) (right-hand and left-hand) of latera, of about twice the size of the two anterior 
ones, which must have come from the carinal half of the whorl, but the exact position of 
which I cannot tell. These latter larger latera are thin, and considerably convex ; they are 
transversely elongated, and, in their longer axis, are rather more than half the length of the 
terga of the same individual. In shape they may be almost compared to one valve of a 
Donax ; being sub-trigonal, with the two upper sides unequal in length, and with the third 
side arched and protuberant at a point about one third of the entire length of the valve 
from the narrower end : to this protuberant point, a well-defined ridge runs from the apex 



POLLICIPES. 67 

of the valve j internally (i), the lines of growth round the upper margins show that the two 
upper sides and the apex overlapped freely the valves on each side of them. This valve, 
as stated in the preliminary remarks, has naturally been always mistaken for a scutum ; 
but the manner in which it has overlapped other valves (as shown by the internal view), on 
both its upper margins, and the entire absence of all hollow for the adductor muscle prove 
that this view is quite erroneous. 

The two rostral or anterior small latera resemble each other (the anterior one being a 
little the smallest), and likewise the largest carinal latera just described ; the external 
ridge, however, here runs to a much more nearly central point of the basal margin ; and 
the shorter and more pointed portion of the basal margin is just perceptibly hollowed out. 
The more pointed end is directed towards the carina. In the first latus this end abuts 
against (no doubt thus producing) the external ridge on the second latus, which it overlaps. 
Internally, under the apex, there is a central crest (/), exactly as in the latera of the lower 
whorl in large specimens of the recent P. mitetta, caused likewise by the overlapping of the 
valves : in the above -described large carinal latera, however, this internal crest is absent, 
showing that the arrangement of these latter valves differed from that of the two anterior 
ones. 

With respect to the number of valves in the whole capitulum, it is almost useless to 
speculate : we have two scuta, two terga, two upper latera, two rostra, and we may, per- 
haps, infer two carina?, making ten valves, we know of three pair of lower latera, making 
sixteen valves : I believe there must have existed some other latera, but probably only 
a few more ; for these valves, especially the carinal pair, are much larger, in proportion to 
the scuta and terga, than in any recent Pollicipes. Probably the lower latera, together with 
the sub-rostrum, and perhaps a sub-carina, formed only a single lower whorl. 

Size : in the individual here described, the carina and terga, were equal in length to 
each other, and "65 of an inch long. In Mr. Bowerbank's collection there is a lower latus 
of nearly double the size of the same valve in this individual, which shows that this species 
attains large dimensions. 

Affinities. As before remarked, this species is very closely related to the cretaceous 
P. glaher, of which it is evidently the representative in the Gault ; the chief difference 
consisting in the more elongated form and greater size of the upper latera, which, in 
P. unguis, exceed half the length of the tergum, whereas in P. glaber they are only one 
third of its length. The carina, in the present species, has its basal margin, perhaps, less 
pointed, and has a narrow linear channel along its edges ; but I am not at all sure that this 
latter character does not vary. Lastly, the anterior lower latera in P. unguis are thinner, 
and rather more convex, with the basal margin more arched and protuberant, with the 
external oblique ridge very much more central. 

Considering the characters of both species taken together, namely, the two ridges pre* 
ceeding from the umbo of the scuta to the baso-lateral angle and basal margin, — the 
triangular shape of the upper latera, — the considerable size of the rostrum, and the ridges 



68 FOSSIL CIRRIPEDIA. 

on the lower latera, — this species comes much nearer to P. mitella than to any other recent 
species ; I believe, however, that it must have had much fewer valves. 1 From the growth 

1 13. Pollicipes validtjs. Tab. IV, fig. 2. 

Pollicipes validtjs. Steenstrup. Kroyer's Tidsskrift, 1839, pi. v, figs. 28 — 32. 

P. scutis crassissimis, angustis ; margine occludente externe costd rotundatd forti firmato ; intus promi- 
nentia rostrali infra marginem rectum basalem dependente ; costd ab apice ad marginem basalem propius 
ad rostralem quam ad basi-lateralem angulum accedente. Carina lavissimd, transverse semicylindricd ; parte 
superiori libere prominente, interne aut plana aut cristd centrali instructd. 

Scuta, extraordinarily thick, narrow ; occludent margin exteriorly strengthened by a rounded, strong 
ridge ; internally, at the rostral angle, a blunt tooth depends beneath the straight basal margin : the ridge 
running from the apex to the basal margin is nearer to the rostral than to the baso-lateral angle. Carina 
very smooth, transversely semi-cylindrical ; upper freely projecting portion internally, either solid and flat, 
or with a central prominent crest. 

Scania, Sweden. Mus. Univers. Copenhagen. Petersberg, near Maestricht. 

My materials consist of several scuta and carinse, sent me by Professor Steenstrup as belonging to the 
same species, which is likewise the opinion of that able collector, M. Angelin, who has found this species in 
various localities in Scania. A very fine carina, from near Maestricht, has been sent to me by Krantz, of Bonn. 

Valves, remarkably thick, massive, and strong. Scuta (fig. 2, e,f, g, nat. size) elongated, being twice 
as long as broad ; slightly convex ; whole upper part bent towards the terga ; surface, in some specimens, 
with traces of longitudinal striae. Basal margin formed obscurely (partly owing to the rubbed condition of 
all the specimens), by two lines meeting each other at a very open angle. Occludent margin much arched, 
forming with the basal margin, taken as a whole, an angle of about 60° : tergo-lateral margin nearly straight, 
forming a rectangle with the adjoining portion of the basal margin. That part of the valve formed by the 
upturned zones of growth is narrow, being, in the widest part, barely half the width of the other part of the 
valve. The occludent margin is strengthened by a rounded, strong, projecting ledge, running along its entire 
length ; the basal end of this ledge, and consequently the lines of growth crossing it, are oblique, and 
slightly upturned. A slight ridge, or angle, runs from the apex to the baso-lateral angle, and a second, still 
slighter ridge, to a point in the basal margin rather nearer to the rostral than to the baso-lateral angle. 
Close to the rostral angle, the internal lamina of the shell is produced downwards into a strong, blunt tooth, 
which (when not too much worn) can be seen from the external side, depending beneath the basal margin : 
this is the most singular character of the species. The internal occludent edge is broad (and of nearly 
the same breadth throughout the whole upper part of the valve), flat, and marked by lines of growth : this 
striated internal edge is separated from the smooth, depending, rostral tooth, by a very oblique line. The 
pit for the adductor muscle is very deep : above this pit there does not appear to have been any furrow on 
either margin, or any marked central prominence. 

Carina (fig. 2, a, b, c, d, nat. size), broad, extremely solid, much bowed inwards ; the upper part, even 
more than half the valve in length, must have freely projected; exterior surface transversely semi-cylindrical, 
or rather steeper than a semi-cylinder, but not at all carinated : basal margin not at all protuberant ; lateral 
angles, or heels, just perceptibly projecting below the central part of the basal margin : in the Maestricht 
specimen it appears that there was a very slight furrow near each exterior lateral margin, making them just 
perceptibly protuberant. The internal, smooth, corium-covered surface is concave, forming almost an equi- 
lateral triangle (b) : above this, the inner freely projecting portion is either filled up flat, or forms a central 
prominent crest (d). 

Affinities. We shall immediately see that this species appears to be most closely allied to P. gracilis, 
of Roemer : it is also allied to P. dorsatus, by the strength of the valves, by the occludent margin of the 



POLLICIPES. 69 

of all the valves, as far as is apparent, being downwards, and from their number, appa- 
rently exceeding 16, I have, without hesitation, ranked this species, (and consequently 
P. glaber,) under the genus Pollicipes. 

scuta being exteriorly strengthened by a prominent ridge, and by the two angular ridges running down to 
the basal margin, and by the general character of the under surface of the apex : it differs widely from P. 
dorsatus, in the rostral, tooth-like process, and in the baso-lateral angle not being widely truncated. The 
carina of these two species also present some points of similarity. In P. validus the centre of the upper 
freely projecting portion often, but not always, has a longitudinal crest or ridge : in the recent P. mitella, 
the degree of filling up of the upper part of the carina varies considerably, and sometimes even a central 
crest is formed in it. 

14. Pollicipes gracilis. Tab. IV, fig. 3. 

Pollicipes gracilis. Roemer. Norddeutsch. Kreidegebirges, 1841, Tab. xvi, fig. 14. 

P. scutis tenuibus, angustis, margine occludente externe costd rotundatd firmato ; intus prominentia 
rostrali infra marginem basalem dependente ; costd ab apice ad marginem basalem centrali : interne^ sulco 
transverso super fossam musculi adductoris, excavato. 

Scuta thin, narrow, occludent margin exteriorly strengthened by a rounded ridge ; internally at the ros- 
tral angle a blunt tooth depends beneath the basal margin ; the ridge running from the apex to the basal 
margin is central. Internally there is a transverse furrow above the hollow for the adductor muscle. 

Unterer Kreidemergel (lower division of Lower Chalk), Hanover. Oberer Kreidemergel, Shaftricht. Mus. Roemer. 

Roemer has most kindly sent me a beautifully preserved scutum, which he has fully described in the above 
quoted work. It is unfortunately rather young, being *6 of an inch in length. It resembles so closely 
the same valve in P. validus, that had it not already received a name I should not have affixed one ; still I 
think it is perhaps a distinct species. It will be quite superfluous to do more than describe the few points 
of difference, of which the chief and most conspicuous one is the much greater thinness of the whole valve. 
I should, however, state that I have not seen any specimen of P. validus so small, the smallest being more 
than one third longer. The second chief difference is that the medial longitudinal ridge here runs to a 
central point of the basal margin instead of to a point nearer to the rostral angle. I think the basal margin 
is more conspicuously formed by two distinct lines, meeting each other at the above point. Internally the 
third and perhaps most important distinction is the presence of a moderately deep and large depression or 
fold, marked by lines of growth, extending inwards from the occludent margin, just above the pit for the 
adductor muscle, and as far inwards at the middle of the pit : this fold precisely resembles that which 
occurs in the same position in Scalpellum magnum, vulgare, and in some Pollicipes, as P. Angelini. The 
rostral tooth appears to have been larger. Owing to the thinness of the valve, the internal occludent edge 
does not here present a flat striated ledge. 

I must again repeat that I doubt whether the differences here pointed out are specific : I have seen nearly 
all such peculiarities variable in other species. I am, however, in some degree strengthened in leaving the 
P. gracilis distinct, by the supposition that it is the representative species in the true Chalk of the P. 
validus of the uppermost stage. 

15. Pollicipes dorsatus. 

Pollicipes dorsatus. Steenstrup. Kroyer's Naturhist. Tidsskrift, 1839, b. ii, h. iv, pi. v, figs. 27 and 30. 

P. scutis crassis ; ad form am trianguli cequianguli accedentibus ; margine occludente externe costd 
rotundatd firmato ; angulo baso-laterali late truncato, dimidium longitudinis marginis basalis veri, cequante ; 



70 FOSSIL CIRMPEDIA. 

ff. Scuta, aid longitudinaliter aut transverse (i. e. secundum tineas incrementi) costata. 
16. Pollicipes striatus. Tab. IV, fig. 5. 

P. valvis longitudinaliter striatis : scutis sub-triangulis, margine ter go -later ali valde 
arcuato et prominente ; costd obscurd, rotundatd, ab apice ad angulum baso-lateralem de- 
cur rente ; interna apicis superficie solide repletd, sulcus manifestus deest. 

Valves longitudinally striated : scuta subtriangular, with the tergo-lateral margin much 

terffo-laterali segmento ex lineis incrementi reflexis formato, angustissimo. Tergorum, costd rectd, lata, pro- 

clivi ab apice ad angulum basalem decurrente ; angulo basali scutum versus, oblique truncato. 

Scuta thick, approaching to an equilateral triangle ; occludent margin exteriorly strengthened by a 

rounded ridge ; haso-lateral angle widely truncated, equalling half the length of the true basal margin ; 

tergo-lateral segment formed by the upturned lines of growth, extremely narrow. Terga with a straight, 

broad, steep-sided ridge running from the apex to the basal angle, which latter on the scutal side is obliquely 

truncated. 

Faxoe ; Denmark. Mus. Univers., Copenhagen. 

Professor Steenstrup has kindly sent for my examination a large series of scuta, terga, and carinse, all 
from the same formation of Faxoe, and no doubt belonging to the same species. 

All the valves are rather strong and thick ; they are smooth, with the exception of the fine lines of growth. 
Scuta (fig. 4/, nat. size, in a reversed position compared with the other figures,) triangular, with the baso- 
lateral angle largely truncated ; considerably convex ; breadth, nearly equalling three fourths of length ; 
apex acuminated, slightly curved towards the terga. Occludent margin nearly straight, forming an angle 
considerably less than a right angle with the basal margin ; this latter is short and nearly straight ; the 
tergo-lateral margin is just perceptibly concave ; a very narrow slip is formed along this margin by upturned 
lines of growth ; this margin, if produced, would meet the basal margin produced, at a less angle than the 
occludent margin does. The baso-lateral angle is so largely truncated that the side so formed is half the 
length of the proper basal margin, which it meets at a very large angle ; it is doubtful whether it should be 
called part of the basal, or of the tergo-lateral margin ; it touches, I suspect, exclusively the latera, but in 
comparison with the other valves of this genus I do not think it corresponds with the ordinary tergo-lateral 
margin. Externally there is a slightly depressed line running from the apex parallel and near to the occlu- 
dent margin, causing this edge of the valve to appear convex and protuberant ; two slight ridges also run 
from the apex — one to the angle uniting the basal and the above-described truncated margin, and the other 
line, which is rather fainter, runs down the valve half way between the first line and the occludent margin. 
Internally, there is a deep, upwardly pointed pit for the adductor muscle ; the internal occludent edge keeps 
the same width up to the apex ; the internal tergal margin has no furrow, but is slightly indented, and must 
have, to a certain extent, overlapped the thin edge of the terga, and thus partly locked the valves together. 
Terga (fig. 4, d, e), rhomboidal, much elongated, almost flat ; occludent and upper carinal margins 
(meeting each other at less than a right angle) much shorter than the lower carinal and scutal margins. 
From the apex to the basal angle, down almost exactly the middle of the valve, a flat-topped ridge, either 
absolutely straight or nearly so, runs, and widens considerably in its downward course ; its carinal side is 
steep, its opposite side less so ; the oblique end of this ridge forms the basal angle, which is nearly, but not 
exactly parallel to the upper carinal margin. There is no depression parallel to the occludent margin. 

Carina (fig. 4, a, b, c) much elongated, very slightly bowed inwards ; transversely, arched flatly ; barely 
sub-carinated ; basal margin almost rectangularly protuberant. Apparently, a full upper half of the carina 
projected freely ; this part is filled up flat and solid (c) ; a rather wide inner margin of the carina extending 



POLLICIPES. 71 

arched and protuberant ; an obscure rounded ridge runs from the apex to the baso-lateral 
angle ; apex with the internal surface filled up solid, without any distinct furrow. 

Upper Cbalk, Norwich. Mus. Fitch, 

My materials consist of two scuta, of which one is young ; and of some terga and a 
carina which I provisionally here describe. 

Scuta (fig. 5, c), broad, moderately convex ; rather thick and strong ; surface strongly 
ribbed from the apex to the basal margin ; ribs rather broad : apex much acuminated ; 
occludent margin nearly straight, at right angles to the basal margin, as is also the lower part 
of the tergo -lateral margin ; the upper part of this latter margin is inflected. Abroad, very 
slightly prominent ridge runs from the apex to the baso-lateral angle, which is broad and 
rounded, but not prominent. There is no second ridge from the apex to the basal margin, 
but along the line where such occurs in P. glaber, the valve is rather abruptly arched ; and 
in the younger specimens, a distinct trace of a ridge can be seen. Internally, the pit for 
the adductor muscle is conspicuous ; the whole upper part of the valve is filled up and 
rendered solid ; the internal occludent edge does not widen above the adductor pit ; on the 
tergal margin a wide internal ledge is marked by lines of growth, is slightly concave, 
irregular, but not furrowed ; on its surface, however, in the larger specimens, there is, 
some little way below the apex, a small ridge and linear hollow, which probably affected, 
in some peculiar way, the shape of the terga; but I doubt whether this structure is 
constant. In general external aspect, this valve comes nearest to the Scalpettum arcuatum ; 
but its tergo-lateral margin not being angularly bent, its baso-lateral angle not being 
prominent, and the greater width of the ridges easily serve to distinguish it. 

Terga (fig. 5, b ; in a reversed position compared to the other figures) . Mr. Sowerby has 
figured a tergum in the ' Mineral Conchology,' Plate 606, fig. 1, under the name of P. sul- 
catus, from the Upper Chalk, and I have had given me by Mr. Woodward a second specimen 
from the Upper Chalk of Norwich. Prom reasons stated under my description of Scatpeilum 
maximum, var. sulcatum, I do not think it can belong to that species ; and from one trifling 
character, namely, the kind of ridge running from the apex to the basal angle, most likely 
it belonged to a Pollicipes ; and as only one species in the Norwich beds is as yet known 
to be ribbed longitudinally, I believe that I have rightly attributed this valve to P. striatus. 

almost to the base, is marked by lines of growth, showing that the valve rather widely overlapped the terga. 
Amongst the specimens there is one very narrow valve, curled a little laterally as well as inwards ; I believe 
it to have belonged to a young and injured or monstrous individual. 

Affinities. The P. dorsatus, though most readily distinguished from the P. elegans of the same Faxoe 
formation, by the absence of ridges on the valves parallel to the lines of growth, and likewise by the absence 
of a strong ridge, running from the apex to the baso-lateral angle of the scuta, certainly has a considerable 
affinity to it, as is particularly manifest in comparing the terga of the two species ; in the scuta of P. elegans 
(fig. 9 c) the truncated end of the oblique wall-sided ridge is obviously analogous with the broad truncated 
baso-lateral angle in P. dorsatus. This species is also related, as we have seen, to P. validus, and P. gracilis ; 
again, also, it is related to the tertiary P. carinatus, and through it remotely to the recent P. mitella. 



72 FOSSIL CIRRIPEDIA. 

Valve much elongated, sub-rhomboidal ; whole surface rather plainly ribbed longitudinally. 
A larger rounded ridge, with the surface of the valve depressed on the carinal side of it, 
runs in a slightly curved line, at about one third of the entire width of the valve from the 
carinal margin, down to the sharp basal angle. Apex narrow, much produced : occludent 
margin about equal in length to the scutal margin : upper and lower carinal margins almost 
running into each other. Parallel to the occludent margin, the valve is depressed, with a 
raised plait in the middle of the depression. Except in the more elongated form, and in 
the character of the ridge running from the apex to the basal angle, this valve is barely 
distinguishable from the tergum of 8. arcuatum (with its varieties) found in the Chalk 
Marl and Gault. 

Carina (fig. 4, a). In Mr. Tennant's collection there is a carina from the Chalk of Kent, 
different from any other seen by me, and which, from being plainly ridged, or rather fur- 
rowed longitudinally, I provisionally describe here. Valve thin, triangular, moderately 
tapering ; very slightly bowed inwards ; transversely, very flatly arched ; plainly sub- 
carinated ; lateral edges narrowly and much inflected ; basal margin rectangularly pointed. 
A very small portion of the valve projected freely. The internal concavity of the valve is 
angular, instead of, as usual, being rounded. The whole exterior surface, except close on 
each side of the central ridge, is longitudinally furrowed. 

17. Pollicipes semilatus. Tab. IV, fig. 6. 

P. valvis longitudinaliter et transverse costatis : scutorum margine basali brevi, recto> 
cum margine ocdudentc unguium rectum formante : costd, parietali, 1 tenuissimd ab apice ad 
unguium prominentem basi-lateralem decurrente ; hac valvam in duas partes inaquules 
dividit, e quibus portio tergo-Jateralis latior est. 

Valves longitudinally and transversely ridged : scuta with the basal margin short, 
straight, forming a rectangle with the occludent margin ; a very narrow wall-sided ridge 
runs from the apex to the prominent baso-lateral angle, and divides the valve into two 
unequal portions, of which the tergo-lateral portion is the broadest. 

Chalk Detritus, Charing, Kent. Mus. Harris. 

General Remarks. I know this species only from one minute broken scutum (*15 of an 
inch in length), with its surface somewhat disintegrated ; but it is certainly distinct from 
the other species hitherto described. The Chalk detritus at Charing is derived from the 
upper and lower Chalk and Chalk Marl. 

Scutum ; the surface is marked by narrow, square-edged, longitudinal ridges, placed 
rather distant from each other ; each zone of growth appears (for the surface is much dis- 
integrated) to have had a prominent plait or ridge which, consequently, runs in lines 
transverse to the longitudinal ridges. The upper part of the valve is only moderately 

1 Parietali, i. e. lateribus utrinque perpendicularibus. 



POLLICIPES. 73 

pointed : the basal margin is short and straight ; it forms, with the occludent margin, a 
rio-ht angle, and with the lower part of the tergo-lateral margin, an angle rather above a 
rio-ht angle. A quite narrow ridge, having perpendicular sides, only about twice as wide as 
the other ridges, runs in a slightly curved course from the apex to the baso-lateral angle ; 
at which angle the ridge, as it appears, must have formed a projection. The tergo-lateral 
portion of the valve, formed by the upturned zones of growth, is unusually broad, rather 
exceeding in width the rest of the valve ; so that the ridge, running from the apex to the 
baso-lateral angle, divides the valve into two only slightly unequal portions, of which the 
tergo-lateral portion is the broadest. Internally, the upper part, above the pit for the 
adductor muscle, is along the middle, slightly prominent • on the tergal side, barely fur- 
rowed, and on the occludent side, slightly hollowed out. This species is allied to the 
three following species, and is, in some respects, intermediate between them and 
P. striatus. 



18. Pollicipes rigidus. Tab. IV, fig. 7. 

Pollicipes rigidus. J . Sowerby. Trans. Geolog. Soc, 2d series, vol. iv, 1836, pi. xi, 
fig. 6* (carina et scuta). 

P. vahis transverse costatis : scutorum margine basali recto, cum margine occludente 
angulum recto majorem formante ; costd angustissimd, parietali, ab apice ad angulum baso- 
lateralem decurrente : tergis, costd curvd, parietali, ad angulum basalem decurrente instructis; 
apice basali in prominentiam parvam terminante, lateribus prominentia par allelis. 

Valves transversely ridged. Scuta with the basal margin straight, forming above a right 
angle with the occludent margin ; a wall-sided, very narrow ridge runs from the apex to 
the baso-lateral angle. Terga with a curved, wall-sided ridge running to the basal angle, 
which latter terminates in a little, parallel-sided projection. 

Gault : Folkstone and Maidstone, Kent. Eastweare Bay, Sussex. Mus. J. Sowerby, Bowerbank, 
Brit. Mus., J. Morris, Flower. 

General remarks. This species appears rather common. I have scuta, terga, and 
carina?, which I infer without hesitation belong to the same species, from the similarity of 
their peculiar surfaces, and from their having been found frequently at the same place and 
in the same formation. 

Description. All the valves have their surfaces conspicuously marked with sharp, nar- 
row, steep-sided prominent plaits parallel to the lines of growth ; each periodical zone of 
growth seems to have been completed by the formation of one of these projecting plaits ; 
the interspaces between the plaits, both on the scuta and carina, are either smooth, or more 
or less plainly fluted with fine longitudinal ridges ; these apparently are in some instances 

k 



74 FOSSIL CIRRIPEDIA. 

only rendered visible or prominent by disintegration ; the transverse plaits are also rendered 
more prominent by disintegration. 

Scutum (Tab. IV, fig. 7, d,f) elongated, triangular, but with the baso-lateral corner 
produced; convex, with the upper part almost semi-conical; apex much pointed, and 
curved towards the carina; occludent margin considerably arched; basal margin short, 
forming, with the occludent margin, an angle considerably above a rectangle, and therefore 
causing the baso-lateral angle to be much produced. The tergo-lateral margin is in the 
upper part slightly concave, in the lower part rounded and protuberant. The baso-lateral 
angle is broad and rounded, but with a small, central, square-sided prominence, formed by 
the projection of the ridge running from this angle to the apex. This ridge is very con- 
spicuous, it is narrow, being not above one third of the width of an average zone of growth, 
increasing very little in width downwards ; it is wall-sided, that is, has its sides absolutely 
perpendicular ; its summit is surmounted by transverse plaits, really continuous (but not 
at first appearing so) with those on the surface of the valve on both sides of it. This ridge 
runs in a curved line, nearly parallel to the occludent margin. Internally (/), the occludent 
edge is broad and flat, and is marked with lines of growth ; it becomes wide at the apex ; 
there is a rather deep furrow close to the tergal margin, but there is no trace of ia central 
ridge ; the pit for the adductor muscle lies quite close under the furrow and flat occludent 
edge ; the nearest approach to this structure of the under side of the apex of the scuta, is 
in the P. glaber. 

Terga (fig. 7, e). On this valve the plaits, in specimens which have not undergone any 
disintegration, are much less strongly pronounced than on the scuta and carina ; in shape, 
sub-rhomboidal, not quite flat ; a curved, wall-sided, narrow ridge (like that on the scutum) 
runs from the pointed, slightly curved apex to the sharp basal angle, and itself projects 
as a little point, with parallel sides ; the growth -plaits extend across this ridge, which 
runs at about one third of the entire width of the valve from the carinal margin. The 
upper carinal margin is very slightly longer than the lower carinal margin, with which it is 
almost blended by a continuous curve. The occludent is shorter than the scutal margin ; 
the edge of the shell close to the former is rounded and protuberant, and parallel to this 
rounded edge the valve is slightly depressed, and correspondingly slightly hollowed out on 
the scutal margin ; internally there are lines of growth along the upper carinal and occlu- 
dent margins. Carina (fig. 7, a, b, c) widening rather slowly from the apex to the base ; 
almost semi-cylindrical ; very slightly bowed inwards ; not in the least carinated ; basal 
margin not at all protuberant. The transverse plaits are, in the three specimens which I 
have seen of the carina, undulatory ; on the very narrow lateral margins (b) the plaits are 
obliquely upturned. The upper part of the carina projected freely ; this part, and both 
margins, are internally marked by lines of growth. 

Dimensions. Most of the specimens are rather small, but I have seen one tergum seven- 
tenths of an inch in length. 

Affinities. This species, with the two following P. fallax and elegans, forma little group 



POLLICIPES. 75 

closely related to each other, and in a far less degree to P. dorsatus. Some remarks on 
their diagnostic characters will be given under the two following species. All three species 
are remarkable by the peculiar form of their scuta, which have so much resemblance (espe- 
cially in P. rigidus) to the terga of other cirripedes, that until I examined their under 
surfaces I was not sure which valves they were. The conspicuous ridges running obliquely 
downwards from the apices of the scuta and terga, I have little doubt were due to the carina 
and rostrum largely overlapping these valves, and to the presence of large upper latera, so 
that the lower angles of the scuta and terga were closely wedged between these valves. 



19. Pollicipes fallax. Tab. IV, fig. 8. 

P. valvis transverse costatis : scutis, margine basalt non recto, angulum pcene rectum 
cum margine occludente formante ; costd, parietibus obliquis, ab apice ad angulum baso-late- 
ralem decurrente .• tergis, costd curvatd, parietibus obliquis, ad angulum basalem latum, 
rotundatum , decurrente. 

Valves transversely ridged. Scuta, with the basal margin not straight, forming nearly 
a right angle with the occludent margin ; a ridge having sloping sides runs from the apex 
to the baso-lateral angle. Terga, with a curved ridge having sloping sides, runs to the 
broad, rounded basal angle. 

Upper Chalk, Norwich. Mus. Fitch. Maastricht Formation, Balsberg and Kopinge, Scania. Mus. 
Univers. Copenhagen. Oberer Kreidemergel (Upper Chalk), Gehrden in Hanover. Mus. Roemer. 

The specimens which I have seen, consist of three right-hand scuta and one tergum, in 
the possession of Mr. Fitch, from Norwich ; of two scuta and two terga collected by M. 
Angelin, in Scania, and forwarded to me by Professor Steenstrup, and of a scutum from 
Hanover, sent to me by Roemer, together with specimens of his P. uncinatus. 

Description. The valves are moderately thick ; they are conspicuously marked with 
rather wide prominent ridges, forming the basal edges of each zone of growth ; they seem 
in both scuta and terga most strongly developed near the occludent margins. 

Scuta (fig. 8, a) almost triangular, moderately convex ; occludent margin considerably 
arched in the upper part, and bowed towards the terga ; basal margin not straight, with 
a short portion close to the rostral angle forming a rectangle with the occludent margin ; 
the remaining portion, if produced, would form a rather larger angle with it ; hence the 
baso-lateral portion of the whole valve is somewhat protuberant. The tergo-lateral margin 
is in the upper part slightly hollowed out, and in the lower part almost straight. A very 
slightly curved ridge runs from the apex to the baso-lateral angle, which is broad, rounded, 
and not prominent ; the ridge has sloping, not wall-like sides. Internally the structure of 
the upper part closely resembles that of P. rigidus. The furrow on the tergal side is rather 



76 FOSSIL CIRRIPEDIA. 

narrower, and a central portion of the inner occludent margin (marked with lines of growth) 
here forms a slightly prominent ridge. In one of the three specimens, the baso-lateral 
portion of the valve was considerably more produced than in the other two. Terga 
(fig. 8, b), rather broad, considerably convex ; apex pointed, and much curled towards the 
scuta ; upper carinal margin unusually arched, slightly longer than the lower carinal margin ; 
there is a deep depression parallel to the occludent margin, which is itself rounded, protube- 
rant, and considerably shorter than the scutal margin. A curved ridge, projecting up above 
the general surface of the valve, with sloping sides, runs from the apex to the basal angle, 
which latter is broad and rounded ; the ridge runs down nearly the middle of the valve. 
Dimensions. Length of longest scutum - 65 of an inch. 1 

1 20. Pollicipes elegans. Tab. IV, fig. 9. 

P. valvis longitudinaliter et transverse striatis : scutoram margine basalt recto, cum margine occludente 
unguium recto paulo major em formante ; costd parietali, latiore quam pro solitd incrementorum latitudine, 
ab apice ad unguium baso-lateralem decurrente : tergorum costd parietali, recta, ad apicem basalem, acumi- 
natum decurrente. 

Valves longitudinally and transversely striated. Scuta with the basal margin straight, forming an angle 
rather above a rectangle with the occludent margin ; a wall-sided ridge, which is broader than the average 
width of the zones of growth, runs from the apex to the baso-lateral angle. Terga with a straight, wall- 
sided ridge running to the pointed basal angle. 

Maestiicht Formation, Faxoe, Denmark. Ignaberga, Scania. 

I am indebted to Professor Steenstrup for an examination of three scuta, a tergum and two carinse, 
firmly embedded in the fragmentary coral-rock of Faxoe, and of two very fine scuta from Scania, collected 
by M. Angelin. The valves have each zone of growth raised into a ridge like the roof of a house ; the 
interspaces between these ridges are marked by longitudinal striae. Length of largest scutum 1*1 of an inch. 

Scuta (fig. 9, c), with the apex acuminated, much curved towards the terga ; occludent margin either slightly 
or much arched ; forming an angle rather above a right angle with the straight basal margin ; tergo-lateral 
margin slightly (but to a variable degree) hollowed out in upper part, and nearly straight in the lower part, 
where it forms nearly a right angle with the basal margin. Baso-lateral angle generally very broad, 
rounded, almost obliquely truncated. From the apex to this angle a gently curved, broad, wall-sided ridge 
runs, which very perceptibly widens in its downward course ; it is generally wider than the average width of 
each zone of growth. Internally the occludent margin is very wide, flat, and marked with ridges; the pit 
for the adductor muscle is deep. 

Terga (fig. 9, d) broad, rhomboidal, slightly convex ; basal angle apparently (for specimen is broken) 
blunt, and obliquely truncated ; upper carinal and occludent margins nearly straight ; shorter than lower 
carinal and scutal margins ; valve considerably depressed in a line parallel with the occludent margin, close 
to which the margin itself is convex and arched. The wall-sided ridge which runs from the apex to the 
basal angle is straight, (not quite correctly represented at d), and widens considerably in its downward 
course : it is not so wide proportionally to the whole valve as the ridge on the scuta, for it is here only a 
little above half as wide as the average width of each zone of growth ; it runs at above one third of the width 
of the entire valve from the carinal margin : the plaits cross its summit in a course not exactly corresponding 
with those on the valve on each side of it ; probably this ridge projected and formed, as in P. rigidus, the 
basal point of the valve. 

Carina (fig. 9, a, b) imperfect, elongated, that is, tapering gradually downwards from the apex ; some- 



POLLICIPES. 77 



Species aliquot quas, scutis incognifis, auctores a valvis insignibus nomindrunt, hie pro 

ten/pore solwnmodo iterum describuntur. 
Species named by authors from remarkable valves, but their scuta being unknown are 

here only provisionally re-described. 

21. Pollicipes bronnii. Tab. IV, fig. 10. 

Pollicipes bronnii. F. Roemer. Verstein. Norddeutschen Kreidegebirges, p. 103, 
Tab. xvi, fig. 8. 

P. carina Icevi, subcarinatd, margine basali arcuato et turgido ; totd valvd vel extrorsimi 
arcuatd vel pane recta ; interne, costis duabus elevatis ad utr unique latus partis superioris 
libere promine?itis. 

Carina smooth, subcarinated ; basal margin arched and protuberant ; whole valve 
either outwardly bowed or nearly straight. Internally, the upper, freely-projecting portion, 
has on each side a prominent ridge. 

Upper Greensand, Warminster, England. Mus. J. Tennant. Hils-conglomerat bei Essen. Mus. Univers., 
Copenhagen. 

My materials consist of some specimens lent me by the kindness of Professor Steen- 
strup ; they consist, as well as those described by Roemer, only of the carina ; their sur- 
faces are considerably worn. The Hils-conglomerat of Essen is considered by Roemer as 
the equivalent of one of the lower beds of the Lower Greensand; but recently MM. 
Saemaan and Geinitz have shown that it really corresponds with the Upper Greensand. 
Professor Tennant has lent me a single broken carina from Warminster, which cannot, in 
our present state of knowledge, be specifically separated from the typical continental 
specimens, though, as we shall see, slightly differing from them. 

I will first describe the foreign specimens. 

Carina (Tab. IV, fig. 10) strong, massive, triangular, about twice as high as broad; but 

what bowed inwards ; basal margin not in the least protuberant ; not carinated, but steeply convex, so that 
a section of the base (b) has steeper sides than a semi-circle. 

Diagnostic characters. In comparing the scuta of P. riyidus, fallax, and elegans, the latter can be at 
once distinguished from P. fallax by the ridge running from the apex having perpendicular or wall-sides, 
and by the straightness of the basal margin ; from P. rigidus, by the ridge being much broader ; P. fallax 
differs from both, in the absence of longitudinal striae. In the terga, P. elegans differs from both P. rigidus 
and fallax, in the ridge running from the apex to the basal angle being straight, and in its greater breadth, 
and likewise in the shortness of the upper carinal margin. The terga of P. fallax differ from the homo- 
logues of the other two species, in the ridge connecting the upper and lower points, not having perpendicular 
sides. 



78 FOSSIL CIRRIPEDIA. 

the breadth varies with respect to the height ; sub-carinated, slightly bowed outwards or 
backwards, and therefore, in the opposite direction to what is usual ; but the amount of 
curvature varies much : roof arched, inflected along the lateral edges ; basal margin 
protuberant, formed of two curved lines meeting each other at an angle of above 90°. 
The lines of growth on the inflected lateral edges, curve slightly downwards (b), and then 
just perceptibly upwards ; thus, no doubt, making the heels or two basal corners slightly 
prominent. Internally, within the lateral edges (having the lines of growth as just described), 
and separated from them by a distinct indentation, there is on each side (b) a narrow ridge, 
widest at about the middle of the upper half of the valve, and marked with longitudinal 
lines of growth ; these internal lateral ridges have evidently been (as seen in the section) 
(d) formed during the filling up and thickening of the upper, solid, outwardly bowed 
part of the carina, which part, no doubt, projected freely. The lower part of the carina (c) 
is deeply concave. Roomer describes the exterior surface of the shell as marked with 
alternate finer and stronger concentric lines, and rarely along the edges alone with longi- 
tudinal lines. 

The specimens from the Upper Greensand of Warminster differ from the foreign 
specimens, in being very slightly bowed inwards instead of outwards, and in having a 
more tapering form ; but it is precisely in these two respects, that the four foreign specimens 
seen by me vary to a considerable extent ; therefore, at present, the English specimens 
must be ranked under P. Bronnii. 

These valves certainly differ considerably from any other known ones : the slight out- 
ward bowing of the carinas from the Hils-conglomerat is their most conspicuous cha- 
racter, and was present in the four specimens seen by me, and I presume, from Roemer's 
description, is general. In recent species, however, the degree of curvature of the carina 
is often variable : in P. sjnnosus, I have seen some specimens with the upper part of the 
carina even outwardly bent, and others with it straight. The manner in which the upper 
part of the valve has been filled up, having two lateral, inwardly projecting ridges, is unlike 
any other species, and is the main specific character : the carinas of Scalpellum solidulum and 
of the var. cglindraceum of 8. maximum make the nearest approach to this structure. From 
such scanty materials I will not pretend to say to what species this is most closely allied. 



22. Pollicipes planulatus. Tab. IV, fig. 11. 

Polltcipes planulatus. J. Morris. Annals of Nat. Hist. vol. xv, 1845, pi. 6, fig. 2. 
— — /. Sowerby. Min. Conch., pi. 647, fig. 2. 

P. tergis subr/iombicis, latis, lavibus, apice basali late truncato, latitudine dimidium lon- 
gitudinis marginis occludentis aquante ; apice basali truncato angulum rectum cum margine 
scutali formante ; parte superiori marginis scutalis cuspidem latam, rotundatam, laviter 
prominentem formante. 



POLLICIPES. 79 

Terga subrhomboidal, broad, smooth, with the basal angle widely truncated, and 
equalling in breadth half the length of the occludent margin ; the basal truncated side 
forms a right angle with the scutal margin : the uppermost part of the scutal margin forms 
a broad, rounded, slightly projecting point. 

Oxford Clay; Christian Malford and Chippenham, Wilts. Mus. J. Morris. 

This species has been named by Mr. Morris, as it appears to me, from the terga alone, 
which are well figured by him and by Mr. Sowerby : these valves (an inspection of two of 
which from Chippenham, I owe to the kindness of Mr. Morris,) are certainly quite distinct, 
in the extent to which the basal angle is truncated, and in the degree of projection of the 
uppermost part of the scutal margin, from any others which I have seen ; but had they not 
been already named, according to the rule followed here, they would have been passed 
over. 

Terga subrhomboidal, or strictly pentagonal, rather broad, very flat and thin ; basal 
angle truncated, extraordinarily broad, equalling half the length of the occludent margin, 
at right angles to the scutal margin ; no single distinct ridge, or furrow, runs from the 
apex to the broad basal angle : upper and lower carinal margins nearly equal in length •. 
occludent and upper carinal margins meet each other at right angles : occludent margin 
rather shorter than the scutal margin ; with the uppermost portion of the latter slightly 
projecting in a blunt, rounded point, which corresponds with a just perceptibly convex slip 
of valve, extending along the occludent margin in the manner so common in this genus. 
Internally, the surface is smooth ; scarcely even the smallest part of the apex could have 
projected freely. 

Largest specimen nearly a quarter of an inch in length, 



{Published species of Lepadida of a doubtful nature, owing to valves, not typical, having 
been named, or from the specimens having been imperfect, or from the shortness of the 
descriptions.] 

Anatifera turgida. Professor Steenstrup (in Kroyer's Tiddskrift, 2 B. (1839), PI. V, figs. 4—5,) has 
thus named some valves from Scania: he states that the scuta resemble in their growth those of Cineras, 
or his Anatifera cretce, that is the Scalpellum ? cretce of this work. Therefore, I conclude that these valves 
cannot properly be ranked under Anatifera or Lepas. 

Anatifera convexa. Roemer, Norddeutsch. Kreidegebirges, Tab. XVI, fig. 7. I do not consider the 
evidence nearly sufficient to place the valve here described in the genus Anatifera ; it might be one of the 
lower Latera of a species allied to Pollicipes glaber or unguis. Pictet (Traite Elementaire de Paleontologie, 
Tom. Ill, p. 438) states that the specimen in question is Aptychus cretaceus. Several vague notices of 
fossil species of Anatifera or Lepas have appeared at various times, but thev hardlv seem to deserve notice. 



80 FOSSIL CIRRIPEDIA. 

Pollicipes angustatus Geinitz (Die Verstein. von Kieslingswalde des Sachsisch-bohmischen, Kreidegebirges, 
1843, Tab. IV, fig. 10) is almost certainly a Scalpellum, and probably a new species ; the carina is narrow, 
bowed inwards, with the basal margin sharply pointed ; tectum carinated, separated from the parietes by a 
line or ridge, which is said to be enclosed between two fine furrows ; this latter seems to be its most 
singular character. I cannot make out whether it has intra-parietes ; only a single view is given : found 
at Strehlen, Saxony, in Planer-Kalke (chalk marl), and, according to the same author, in his ' Das Quader- 
sandsteingebirge" (p. 100) ; likewise in Greensand, at Essen, in Westphalia. 

The Pollicipes Icevis of J. Sowerby, figured in 'Geological Transactions,' 2d series (1836), Vol. IV. PL 
XI, fig. 5, I consider the same as his P. unguis, fig. 5* ; but the P. Icevis, from Blackdown, PI. XVI, fig. 1, 
seems to be certainly a distinct species, and possibly a Scalpellum: no details are given. The P. radiatus 
of the same author, of the lower greensand (PL XI, fig. 6), is unknown to me ; the tergum figured is like 
that of S. arcuatum ; the upper valve, if a scutum, is very remarkable. 

The name Pollicipes radiatus has been a favorite with authors ; there is a tertiary species so called, 
but not described in 'Mliller's Jahrbuch,' 1835. Koch and Dunker, also, (Norddeutsch. Oolithgebildes, 
1837,) have given this name to a tergum found, according to Roemer, in the Hilsthon or Lower Greensand ; 
I conceive it to be scarcely possible, and very inconvenient, to attach names to terga. 

Michelotti has called, without any description (' Bulletin, Geolog. Societe,' torn. 10, p. 140), a tertiary 
Pollicipes from near Turin, P. antiquus, — a most infelicitous name for a miocene species. 

Roemer in his ' Die Versteinerungen des Norddeutsch. Kreidegebirges,' 1841, has figured, PL 16, fig. 13, 
the above-mentioned P. radiatus, of Koch and Dunker ; also, Pollicipes asper (PL 16, fig. 15), which is 
founded on a fragment of a carina, which I have been permitted to inspect by the great kindness of Roemer; 
it is not sufficiently perfect in the upper part to show whether it probably belonged to Scalpellum or Polli- 
cipes, but apparently to the latter genus ; it is almost certainly distinct from anything which I have seen ; 
the tectum is very flatly arched, and the basal margin (as inferred from the lines of growth) scarcely at all 
protuberant ; it was very slightly bowed inwards ; its surface is covered with faint longitudinal ridges, and 
these are crossed by concentric, leaf-like, projecting lines of growth. The longitudinal ridges cause this valve 
to resemble that of Scalpellum solidulum from Scania, but the basal margin is much less protuberant, the 
tectum more steeply arched, and the whole valve thicker than in that species. The P. asper comes from the 
"Oberer Kreidemergel" (Upper Chalk) of Quedlinburg, in Westphalia. 

Roemer has also described in the same work (PL XVI, fig. 10, a, b, c,) Pollicipes uncinatus from the 
Upper Chalk : the carina (b) has its roof sharply carinated ; its parietes are quite flat, and I presume 
rectangularly inflected, which makes me suppose it may have belonged to a Scalpellum ; basal margin rectan- 
gularly pointed. The valve, described as a tergum (c) I have had, through the kindness of Roemer, an 
opportunity of examining ; it cannot, I conceive, be a tergum ; it is unknown to me ; it appears to 
be a carinal latus ; if so, the valve ought to have been placed in the figure transversely to its present 
position. 

Reuss in Die Verstein.' der Bohmischen Kreideformation (1845) has figured (Tab. V, fig. 43) a carina, 
under the name of Pollicipes conicus, from the Chalk ; I do not consider it to be recognisable. The P. glaber 
in this same work (Tab. V, fig. 45-49, and Tab. XIII, fig. 86 — 91) appears to consist of several species ; on 
the other hand, P. radiatus (Tab. V, fig. 42) appears to be a scutum of the true P. glaber. 

Dr. W. Dunker (Palseontographica : Beitrage zur Naturgeschichte, &c, der Vorwelt von Dr. W. Dunker, 



LORICULA. 81 

and H. Von Meyer, 1848, 1 Band, p. 180, Tab. XXV, fig. 14) has described and figured a valve of Pollicipes 
liasinus : he considers it as one of the great hinder lateral valves — that is, a tergum. This eminent palaeon- 
tologist is well acquainted with the cirripedia, and is therefore probably right in his determination ; but I am 
forced to say that the manner of growth seems to me, if I rightly understand the figure, unlike that of the 
terga in any known Pollicipes. 

Miiller (Aachen, p. 43, Tab. II, fig. 16,) has described Pollicipes ornatissimus ; the valve figured is a 
carina, I believe, of Scalpellum maximum. 

Pollicipes . /. Sowerbij. Geolog. Transact. 2d series, vol. v, pi. ix, fig. 2. 

The specimens here figured by Mr. Sowerby, were found by Mr. Wetherell in a deep well in the London 
Clay, at Hampstead, together with portions of Scalpellum quadratum. The specimens are in a broken con- 
dition, and have heen kindly lent me by Mr. Wetherell : they consist of some fragments of terga, and I 
believe of the basal portion of a scutum, of a carina, and of several latera. The species appears to have 
come near to the Eocene P. reflexus ; but although thinking it distinct, I dare not, considering the imperfect 
state of the scutum, name it. The valves appear to have been much thicker and more rugged than those of 
P. reflexus ; the basal angle of the terga sharper, and the rostral angle of the scutum more abruptly trun- 
cated. The latera are remarkable, and unlike anything which I have seen ; they present several shapes; 
they are all more or less triangular, and their lateral edges are more or less distinctly bordered exteriorly by 
prominent ridges ; one has a central exterior ridge, and its basal margin is protuberant ; another has one of 
its sides at right angles to the basal margin, and the other side largely convex. Not having seen the lower 
latera of the P. reflexus, or indeed of any fossil Pollicipes except of P. glaber and unguis, I cannot use 
characters drawn from the latera as diagnostic. The specimens are extremely small. 



Genus — Loricula. 

Lokicula pulchella. Tab. V, figs. 1 — 4. 

Loricula pulchella. G. B. Sowerby, jun. Annals of Nat. History, vol. xii, 1843, 

p. 260. 

L. capitulo decern (fortasse) valvis instructo. Pedunculo seriebus decern squamarum 
lamum calcarearum instructo ; sex lateralibus milium transverse elongatis ; quatuor termi- 
nalibus angustis ; secundum pedunculi margines rostralem et carinalem decurrit sutura 
medialis recta, squamis non inter secantibus. 

Capitulum with (perhaps) ten valves. Peduncle with ten rows of smooth calcareous 
scales, of which the six lateral rows are much elongated transversely, and the four end 
rows narrow ; along the rostral and carinal margins of the peduncle there is a straight 
medial suture, with the scales not intersecting each other. 

Lower Chalk, Cuxton, near Rochester, Kent. Mus. Wetherell. 

I owe to the kindness of Mr. Wetherell an examination of this beautiful and unique 
specimen, well described and figured by Mr. G. B. Sowerby, Junr., in the ' Annals of 

1 



82 FOSSIL CIRRIPEDIA. 

Natural History.' ' I believe the specimen to be extremely incomplete ; certainly only 
one side of the capitulum and peduncle is preserved ; but on this subject I shall offer a few- 
remarks after describing the specimen in its present state. The specimen was embedded 
outside the cast of an Ammonite : Mr. J. Morris informs me, that he has no doubt that 
the pit at Cuxton is worked in the Lower Chalk. 

The Capitulum (Plate V, fig. 1, of nat. size,) consists of three left-hand valves, which are 
small compared to the size and width of the peduncle, making me believe that the greater 
part of the animal's body was lodged in the peduncle, as in the recent genera, Lithotrya 
and Ibla. Of the three valves, I believe from the general shape and direction of the lines 
of growth, that one (on the right-hand side of the figure) is the scutum ; of the other two, 
I will at present call that nearest to the scutum the first latus, and the other the second 
latus. The three valves are of nearly equal size : they are very slightly convex, quite 
smooth, moderately thick, with the lines of growth fine and obscure. 

Scutum (fig. 3) triangular, apex somewhat produced, and bowed over towards the 
latera : occludent margin slightly arched, forming an angle much less than a rectangle 
with the basal margin, which latter is at right angles with the margin, called in Pollicipes 
tergo-lateral. The lines of growth, in the lower part of the valve, are parallel to the tergo- 
lateral and basal margins ; and as far as I could distinguish in the imperfect condition of 
of the valve, the valve is added to a little during growth at the apex, so that the umbo 
is not situated at the uppermost point. In this important respect Loricula apparently 
resembles the Scalpettum magnum, tuberculatum and creta. 

The first (so-called) latus is nearly flat, triangular, with its apex on a level with that 
of the scutum ; the scutal margin (lying close and rather over the edge of the scutum) 
is just perceptibly arched, and a very little longer than the other lateral margin : the basal 
margin is shorter than the two lateral margins ; growth downwards. 

The second (so called) latus (fig. 2) is triangular, of nearly the same shape with the last 
valve, with the inner basal angle a little produced ; the outer margin is very slightly arched, 
and is shorter than the inner margin ; the main growth is downwards, but the lines curl a 
little up and round the inner margin. The apex of the valve in its present position stands 
a little above the apices of the other two valves, and a wedge-formed open space separates 
the upper part of the two latera. 

Peduncle : this has a most singular, elegant, loricated structure : it is wider than the 
capitulum in the present condition of the latter, and about four times as long. It is 
completely protected by large, smooth, calcareous scales, of which five rows are preserved ; 
and I conceive there can be no doubt from the shape of the end rows, that there originally 
existed a corresponding series on the opposite side, making altogether ten rows. The 
base is sharply pointed, down to which the full complement of scales extends. In each row 

1 Since Mr. Sowerby made his description, Mr. Wetherell has cleared away more of the Chalk, thus 
exposing a row of small scales at the carinal end of the peduncle, not seen by Mr. Sowerby. 



LORICULA. 83 

there are about twenty-one scales, their numbers obviously depending on the age and size 
of the individual. There is one more scale under the second latus (fig. 4, restored figure), 
than under the first latus, and one more under the first latus than under the scutum ; hence 
the summit of the peduncle is obliquely truncated, being lowest at the rostral end : in this 
respect there is some resemblance to the genus Lithotrya. The scales in the three lateral 
rows resemble each other pretty closely in outline : they are transversely elongated, and 
are together about as wide as the three valves of the capitulum ; they are much longer 
than the scales of either end row. The first row of these elongated scales lies directly 
under the scutum, and the other two under the two latera ; so that the lines of junction of 
the three rows of peduncular scales, and of the three valves of the capitulum, correspond. 
The scales in the middle row are rather longer than those on either side of it, are pointed 
at both ends, and have their upper margin very flatly arched and almost square-edged. 
The scales in the row under the second latus are rather wider than it, projecting (which is 
important) beyond its outer edge ; their upper margins are square-edged ; their outer ends 
blunt and truncated; their inner ends pointed. The scales of the third row under the 
scutum are rather less wide than it, and do not reach so far as its outer or rostral angle ; 
their upper margins are arched ; their outer ends bluntly rounded, and their inner ends 
pointed. 

The two end (that is, the rostral and carinal) rows of narrow scales remain to be 
described : those at the rostral end (fig. 3) are as high as the larger lateral scales, but only 
about one fourth as wide : in shape they are almost a rectangular oblong, with their 
upper ends a little rounded, and the outer (with respect to the longitudinal axis of the 
animal) basal angle a little produced ; hence the two lateral margins of the scales of 
this rostral row do not quite match each other ; consequently, to make the animal symme- 
trical, there must have been a corresponding approximate row of small scales on the other 
side of the medial line. The straight inner (both sides of the peduncle being supposed 
to be present) edges of the scales of the row just described, extend rather beyond the 
occludent margin of the scuta. The scales in the carinal row, at the opposite end of the 
peduncle, are not above half the width of those of the rostral row : they are of nearly 
the same shape, but their upper ends are more pointed, and their outer (with respect to 
the medial longitudinal axis of the animal) basal angles more produced ; their straight 
inner margins projected considerably beyond the carinal edge of the second latera : it is 
more obvious in this case than in the rostral row, that there must have been a second 
adjoining row of small scales on the other side of the carinal medial line. The small 
scales of the carinal and rostral rows differ from the others in their inner (that is, close to 
the medial axis), basal angles, not crossing each other ; so that the peduncle could have 
been divided in a medial rostro-carinal plane, without cutting through a single scale. 
Their outer basal angles, on the other hand, intersect the ends of the adjoining large 
lateral scales, like these latter intersect each other, the lines of intersection between them 
being straight and corresponding with the junctions (as already stated) of the scuta and 



84 FOSSIL CIRRIPEDIA. 

latera. The consequence of this arrangement is, that in the alternate whorls there are 
in one, four of the large lateral scales and none of the smaller end scales ; and in the 
whorl, both above and below, only two of these large lateral scales, and four of the carinal 
and rostral scales ; so that the alternate whorls (fig. 4) had an unequal number of scales, 
namely, four and six. 

Growth. — New scales for the peduncle are formed round its upper edge, at the bases 
of the valves of the capitulum, the chief growth of which, as we have seen, is downwards : 
hence, we here have, as in other pedunculated Cirripedia, a principal line of growth round 
the summit of the peduncle. It can be seen that a new scale is first formed under the 
second latus, at the carinal end of the peduncle ; and this agrees with the fact that there 
is one more scale in this row than in that next to it ; and one more in the latter, than in 
the row under the scutum. I may mention, as in conformity with this fact, that in the 
development of the young of Scalpettum vulgare from the larval condition, the calcareous 
scales on the peduncle first appear under the carina. 

Attachment. — With respect to the attachment, Mr. G. B. Sowerby seems to have felt 
great difficulty on account of the peduncle ending in so fine a point ; but the peduncle 
of Scalpettum vulgare, when carefully dissected from the coralline to which it is attached, 
often ends in a much finer point, and is symmetrically attached to the branch by its 
narrow rostral margin. In Loricula, the attachment was probably by one lateral face of 
the lower part of the peduncle ; for it is by no means unusual for the cement-stuff (even 
when proceeding only from the two original central orifices, where the prehensile antennas 
of the larva may still be found) to encroach largely on the peduncle, and thus fix it down. 
The calcareous scales of Pollicipes, and the horny spines of Ibla, may often be found thus 
embedded and firmly fixed to the supporting rock ; it is moreover possible, that in Loricula 
the cement was poured out of orifices, specially situated on one side of the peduncle, as 
takes place along the rostral margin in Scalpellum vulgare, and high up on both sides of 
the peduncle in Lepas fascicular is. 

Dimensions. — Entire length of the specimen rather above one inch : width of widest 
part of the peduncle "G of an inch. 

On the probable condition of the Specimen when perfect. — I have already remarked 
that the three valves of the capitulum, and the five rows of scales on the peduncle, must 
have had an opposite series. I at first thought it extremely improbable that the animal 
should have been split so exactly down the middle ; and I conceive, in most genera, as in the 
common Lepas, it would be quite impossible to effect this ; but in removing the terga and 
scuta from one side of several specimens in Ibla and Lithotrya, it was difficult to prevent 
exactly half of the membrane of the peduncle (which in these genera is thin) being torn 
off with them. Mr. G. B. Sowerby, jun., also has remarked (Annals of Nat. Hist., 1843, 
p. 261), that, owing to the rostral and carinal rows of small scales not intersecting each 
other, the splitting of the specimen into halves along this plane would be much favoured. 
The inner edge of the so-called second latus could not have touched the inner edge of its 



LORICULA. 85 

fellow valve on the opposite side, inasmuch as two small peduncular scales were inserted 
between the ends of the larger peduncular scales, which of themselves projected beyond 
the edges of the second latera : to fill up this hiatus, I conceive there must have once 
existed a carina. For the same reason, I believe, but less strongly, that there also 
existed a rostrum. It may be observed that in the present condition of the specimen, 
the straight lines of intersection between the two outer large peduncular scales and the 
small rostral and carinal scales, do not correspond with junctions of any valves in the 
capituluni; but if, as in our imaginary figure (4), a carina and rostrum be added, 
the above two lines of intersection will correspond with junctions of valves in the capi- 
tuluni; making altogether, on each side, in the capitulum four lines of junction, and in 
the peduncle four lines of intersection. 

At present there is a wedge-formed open space between the first and second latera ; 
and judging from the lifted up position (fig. 1) of the upper scales of the peduncle under 
the second latus, this interspace must originally have been a little larger. In order to com- 
plete the sack to receive and protect the animal's body, this interspace must have been 
closed either by a valve or by a membrane ; — the latter supposition seems to me very 
improbable, considering the closely loricated condition of the whole exterior ; consequently, 
I believe that on each side, a triangular valve with its apex downwards was wedged in 
between the two latera, and that these valves answered to the terga of other Cirripedes. 
On any other view, considering the high improbability of the entire abortion of the terga, 
we must conclude that either the first or second latus was a tergum, though of a totally 
different shape from that valve in every other pedunculated Cirripede ; the other latus, 
moreover, would on such a view be a complete anomaly. 

In the imaginary restored figure (fig. 4), the tergum has its normal shape and manner 
of growth. The first latus now answers to the upper latus in Scalpellum, but it is inter- 
posed to a quite unprecedented extent between the scutum and tergum. The second latus 
is on this view the carinal latus ; and the rostral latus, always smaller than the carinal 
latus, and in S. quadratum and Peronii reduced to a very small size, is here quite aborted. 
A restoration, however, of this nature must always be very doubtful. Finally, I imagine 
that the Loricula was attached by the side of the lower part of its peduncle to some large 
shell or coral, and that a crab or some other animal tore off the upper side, and this subse- 
quently being dropped, became embedded in the chalk ; the corresponding valves with the 
addition of a carina, rostrum, and a pair of terga being left adhering to the surface of 
attachment. 

Affinities. — In an animal so imperfectly known, it is useless to enter on this subject : 
I will only remark that the restored figure comes nearest to the genus Scalpellum ; the 
recent S. ornatum has the scales on the peduncle even proportionally more elongated than 
in Loricula. The chief distinctive character of the genus, as at present imperfectly known, 
lies in the arrangement of the scales on the peduncle. In those species of Pollicipes, Scal- 
pellum, and Lithotrya, in which the scales on the peduncle are symmetrically arranged, 



86 FOSSIL CIRRIPEDIA. 

each scale lies exactly between two scales in the whorls, both above and below : this is, in 
fact, the case with the large lateral scales in Loricula, but the ends of the scales in the 
same whorl, instead of, as is usual, quite or nearly touching each other, are here far 
removed from each other, so that each whorl is broken by wide open spaces. In the 
marked difference in size between the lateral scales and those in the two end rows ; — in 
the latter scales not intersecting each other, but presenting a straight, medial, rostral and 
carinal suture ; — and lastly, in each alternate whorl having a different number of scales, 
namely, four in one and six in the other, Loricula differs from every other known 
Cirripede. 



INDEX. 



[N.B. — The names in italics are either synonyms or doubtful species.] 



Alepas squalicola .... 
Anatifa cancellata 

„ convexa .... 
Anatifera cretce .... 

„ turgida 
Angelin, M., on fossils of Scania . 
Antennae prehensile of larva . 
Aptychus ..... 
Attachment, manner of, in Cirripedes 
Balanus carbonarius 
Calentica Homii .... 
Carina, manner of growth of . 
Clisia or Creusia, fossil 
Corals, fossil, not inhabited by Cirripedes 
Crustacea compared with Cirripedes 
Dana, J. D., on the peduncle of Cirripedes 
Distribution of recent Cirripedes . 

„ fossil Cirripedes 

D'Orbigny, Alcide, on Aptychus 
Fitch, Mr., specimens collected by 

Forchhammer, on the age of the beds 
Scania ...... 

Gray, Mr. J. E., on the genus Scalpellum 

Homologies of the peduncle and capitulum 

Ibla, sex of . 

Leach, Dr., on the genus Scalpellum 

Lepadidae ..... 

Loricula pulchella 

Loven, on the homologies of Cirripedia 

Male Cirripedes .... 

Meyer, on Aptychus 

Morren, on fossil Tubicinella 

Nomenclature of valves 

Nomenclature, rules of . 

Oken, on Mitella .... 

Oxynaspis (genus recent) 

Peduncle, homologies of 

Petzholdt, on Balanus carbonarius 

Pictet, on fossil Crustacea 



Page 
13 
7 
79 
45 
79 
6 

13 

3 

13 

5 

14 

19 

5 

5 

13 

12 

6 

8 

3 

6 



of 



7 

3, 14 
13 
16 
3, 14 
12 
81 
13 
23 
4 
5 
9 
47 
47 
45 
13 
5 



Pollicipes acuminatus 
Angelini 

„ angustatus 

,, an ti quits 

„ asper 

„ Bronnii 

,, carinatus 

,, concinnus 

,, conicus . 

„ dorsatus 

„ elegans 

,, elongatus 

„ fallax 

„ Genus of 

„ glaber . 

,, gracilis 

„ Hausmanni 

,, Icevis 

„ liasinus 

,, medius . 

,, maximus 

„ Nilssonii 

„ ooliticus 

,, ornatissimus 

,, planulatus 

,, politus . 

„ quadricarinatus 

„ reflexus 

„ radiatus 

„ rigidus . 

„ semilatus 

,, solidulus 

„ striatus 

„ Sub-genera of 

,, sulcatus 

„ uncinatus 

„ undulatus 
unguis . 

„ validus . 

„ villosus . 



Page 

. 56 

. 56 

. 80 

. 80 

. 80 

. 77 

. 60 

. 50 

. 80 

. 69 

. 76 

. 55 

. 75 

. 47 

. 61 

. 69 

. 53 
55, 65, 80 

. 81 

. 26 
24, 26, 34 

. 52 

. 51 

. 81 

. 78 

. 54 

. 38 
58, 81 
40, 80 

. 73 

. 72 

. 42 

. 70 
3, 48 

. 26 

. 80 

. 42 

. 64 

. 68 

. 14 



88 



INDEX. 









Page 


Roemer, work of . 


1 


Eules of British Association . 


. 47 


Scalpellum angustum 


. 37 




, arcuatum 


. '0 




, cretaceous species allied to S. r 


nagnum 21 




, (?) cretse 


. 45 




, fossula 


. 24 




, Genus of 


. 13 




, hastatura 


. 37 




, lineatum 


. 35 




, magnum 


. 18 




, maximum 


. 2G 




, maximum, var. cylindraceum 


. 33 




, maximum, var. sulcatum 


. 34 




, ornatum 


15, 23 




, quadratum . 




. 22 




, quadri carina turn . 




. 38 




, rutilum 




15, 23 




, semiporcatum 




. 44 




, simplex 




. 39 




, solidulum 




. 42 




, Sub-genera of 




3, 15 




, trilineatum 




. 38 



Scalpellum tubercul: nm 

,, villosum 

„ vulgare, sex of 

Sexes, peculiarities of, in Scalpellum 
Sowerby, G. B., on a fossil Anatifa 
Sowerby, J., work of 

„ on the genus Xiphidium 

Steenstrup, Professor, work of 

,, „ on age of beds of 

„ „ on P. solidulus 

Stomapoda ..... 
Stutchbury, Mr., on fossil Cirripedes 
Thaliella ... 
Trigonellites 
Tubicinella fossil 
Valves, nomenclature of 
Verruca fossil in the cb*lk 
Wetherell, Mr., on a well at Hampstead 
Xiphidium, Genus 

„ angustum 
„ quadratum . 

,, maximum 



Scania 



Page 

43 

14 

16 

5, 23 

7 

1 

15 

1 

7 

42 

12 

5 

15 

3 

5 

9 

5 

22 

15 

37 

22 

61 



''•it 



TAB. I. 

[In evert/ case riyht-hand Scuta and Terga are figured ; hence the occludent margins are always to the left-hand.'] 

Fig. 1. Scalpelliun magnum : — fig. (a) natural size, the rest magnified twice. 

(a) Imaginary restored figure, of natural size. (h) Rostral latus, inside view of, broad variety. 

(b) Carina, dorsal view. (i) Ditto, outside view of, narrow variety. 

(c) Scutum. (k) Ditto, seen in profile. 

(d) Tergum. (/) Carinal latus, outside view. 

(e) Upper latus. (m) Ditto, inside view. 

(/) Carina, lateral view. (n) Ditto, outside view, broad variety. 

(g) Rostral latus, narrow variety. 

Fig. 2. Scalpellum angustum : — carina, dorsal and lateral views of, copied from Dixon's 
' Geology and Fossils of Sussex,' Tab. xxviii, fig. 9. 

Fig. 3. Scalpellum quadratum, fig. {a) natural size, the rest magnified twice. 

(a) Specimen as found embedded, with the (d) Carina, lateral view. 

valves in nearly their natural positions ; (e) Upper latus. 

the end of the rostral latus ought to (/) Rostral latus. 

touch the short basal side of the upper (g) Carinal latus. 

latus. (A) Scutum, internal view. 

(b) Scutum. (i) Carina, dorsal view. 

(c) Tergum. (k) Ditto, section across middle of valve. 

Fig. 4. Scalpellum fossula; all the figures magnified twice, except (It), which is four 
times the natural size. 

(a) Scutum. (e) Carinal latus. 

(b) Tergum. (/) Scutum, internal view. 

(c) Carina, side view. (g) Carina, dorsal view. 

(d) Upper latus. (h) Ditto, section across middle. 

Fig. 5. Scalpellum trilineatum ; all the figures magnified twice, except (c and d), which 
are four times the natural size. 

(«) Carina, dorsal view. (d) Carina, section of, near apex. 

(b) Ditto, lateral view. (e) Tergum. 

(c) Ditto, section of, lower part. 

Fig. 6. Scalpellum semiporcatum ; scutum, magnified three times. 

Fig. 7. Scalpellum arcuatum; all the figures magnified twice, except {</), which is four 
times the natural size. 

(a) Carina, dorsal view. (e) Surface of carina, much magnified. 

(4) Ditto, lateral view. (/) Scutum. 

(c) Tergum, inside view. (g) Section across carina. 

(</) Ditto, outside view. 

Fig. 8. Scalpellum solidulum ; natural size, except (/). 

(a) Tergum. (d) Carina, section of upper part. 

(b) Carina, doi-sal view. (e) Carinal latus, natural size. 

(c) Ditto, internal, almost lateral, view. (/) Ditto, much magnified. 

Fig. 9. Scalpellum simplex ; twice natural size. 

(a) Carina, dorsal view. (jb) Carina, lateral view. (c) Carina, section of. 

Fig. 10. Scalpellum tuberculatum ; largely magnified. 

(a) Tergum. (d) Scutum ; (e) basal margin, (/) occludent mar- 

(6) Carina, dorsal view. gin ; these two margins ought to form a 

(c) Ditto, lateral view. rather larger angle. 

Fig. 11. Scalpellum (?) cretse; largely magnified. 

(a) Tergum. (b) Carina. (c) Scutum. 



Tab. I 




J.De (' . Sowerty fecit 



* 



TAB. II. 

[/« every case right-hand Scuta and Terga are figured ; hence the occludent margins are always to the left-hand.] 

Fig. 1. Scalpellum maximum, var. typicum. 

(a) Carina, twice natural size. (d) Section of carina across middle of valve, twice 

(b) Ditto, lateral view, natural size. natural size. 

(e) Ditto, ditto, twice natural size. (e) Section of carina across lower part of valve. 

Fig. 2. Scalpellum maximum, var. cylindraceum ; all the figures twice the natural size, 
except the sections. 

(a) Carina. (c) Section of carina, upper part. 

(b) Ditto, lateral view. (d) Ditto, ditto, lower part. 

Fig. 3. Scalpellum maximum, var. sulcatum. 

(a) Carina, twice natural size. (d) Section across the middle of carina, four times 

(b) Ditto, natural size. natural size. 

(c) Ditto, lateral view. 

Fig. 4. Scalpellum maximum ; carinal latus, two varieties. 

(a) Natural size. . (c) Natural size. 

(6) Magnified four times. (d) Magnified four times. 

Fig. 5. Scalpellum maximum; Tergum, Var. I., natural size and twice magnified. 

Fig. 6. Scalpellum maximum ; Tergum, Var. II., natural size and twice magnified. 

Fig. 7. Scalpellum maximum ; Tergum, Var. III. 

(a and a) Natural size, and twice magnified. (b) Internal view, twice magnified. 

Fig. S. Scalpellum maximum; Var. I. 

(a) Scutum, twice natural size : some longi- (b) Scutum, natural size. 

tudinal lines have heen erroneously (c) Ditto, internal view, twice natural size, 
introduced in this engraving. 

Fig. 9. Scalpellum maximum ; Var. II. 

(a) Scutum, outside view, twice natural size. (b) Scutum, internal view, twice natural size, 
(c) Scutum, outside view, natural size. 

Fig. 10. Scalpellum maximum; Var. III. Scutum, inside view, natural size. 

Fig. 11. Scalpellum lineatum. 

(a) Scutum, natural size. (b) Scutum, twice natural size. 

Fig. 12. Scalpellum lineatum ; Carina twice natural size. 

Fig. 13. Scalpellum hastatum. 

(a) Carina, twice natural size. (c) Carina, lateral view, twice natural size. 

(6) Ditto, natural size, lateral view. (d) Ditto, section across middle of valve. 



Tab.U. 




•T.De C.Sowerby feci! . 



TAB. III. 



[/re every case right-hand Scuta and Terga are figured ; hence the occlu ent margins are always to the left-hand. J 

Fig. 1. Pollicipes concinnus; copied from the Mineral Conchology, PL 647. 



(a) Group of specimens as found adhering to 
an Ammonite, of the natural size. 

Fig. 2. Pollicipes ooliticus. 

(a) Carina. 

(b) Rostrum. 

Fig. 3. Pollicipes Hausmanni. 

(a) Carina. 
(6) Scutum. 

Fig. 4. Pollicipes politus. 



(b) Capitulum enlarged. 

(c) Scales of the peduncle magnified. 



(c) Tergum. 

(d) Scutum. 



(c) Scutum, inside view of. 

(d) Tergum. 



Fig. 



(a) Scutum. (b) Small portion of the occludent margin, 

muck magnified, to show the narrow prominent ledge. 

5. Pollicipes elongatus. 

(«) Tergum, about half natural size. (c) Scutum, natural size. 

(b) Scutum, much magnified ; the impression on the chalk gives the general outline. 



Fig. 6. Pollicipes acuminatus. 

(a) Scutum, outside view, figure restored. 

Fig. 7. Pollicipes Angeliui. 

(a) Scutum. 

(6) Ditto, inside view of. 

(e) Tergum. 

Fig. 8. Pollicipes reflexus. 

(a) Carina, dorsal view. 

(b) Ditto, lateral view. 

(c) Ditto, section beneath the middle. 

(d) Tergum. 

Fig. 9. Pollicipes carinatus. 

(a) Tergum. 

(b) Ditto, internal view. 

(c) Carina. 

(d) Scutum. 

(e) Scutum, inside view of. 

Fig. 10. Pollicipes glaber. 

(a) Tergum, broken carina, upper latus, and 

a latus of the lower whorl ; all these 
valves belonged to the same individual, 
and show their relative natural sizes. 
(6) Scutum, natural size, of another indi- 
vidual. 

(c) Ditto, ditto, four times magnified. 

(d) Ditto, inside view of upper part ; the left- 

hand is the occludent margin. 

Fig. 11. Pollicipes Nilssonii. 

(«) Scutum, from a small worn specimen. 

(b) Carina, natural size, lateral view of. 

(c) Ditto, section of, across the middle. 



(b) Inside view of actual specimen. 



(d) Section across the middle of the tergum, to 
show form of surface. 



(e) Scutum ; the letter (e) stands close to the occlu- 
dent margin. 
(/) Upper latus. 



(/) Carina, section of, near apex. 
((/) Rostrum, inside view. 
(A) Ditto, lateral view. 
(t) Ditto, dorsal view. 



(e) Tergum, four limes magnified. 

(/") Carina. 

((/) Ditto, small portion of lateral margin, close 

above the basal margin, much enlarged. 
(h) Rostrum, natural size. 
(i) Ditto, four times magnified. 
(k) Upper latus, three times magnified. 
(I) Latus (probably from near the rostrum) of the 

lower whorl, three times magnified. 



(d) Sub-carina, inside view, natural size. 

(e) Rostrum, inside view, natural size. 



Tab III 




,1 De (' . Sowerhy fecii 



TAB. IV. 



[N.B. — A left-hand scutum (fig. 4,/), and a left-hand tergurn (fig. 5, b), have been accidentally figured instead of right- 
hand valves, as in all the other Figures and Plates ; hence these two hold reversed positions, and in both^ the occludent margins 
are to the right, instead of to the left hand.'] 

Fig. 1. Pollicipes unguis; all the figures except (d) are from the same individual, and 
are magnified twice : (d) is of the natural size. 

(a) Carina. (h) Latus of the lower whorl, probably adjoining 

(6) Tergura, inside view. the carina. 

(c) Ditto, outside view. (i) Ditto, inside view. 

(d) Ditto, ditto, (var., natural size.) (£) Latera of the lower whorl, probably the two 

(e) Rostrum. adjoining the rostrum. 

(f) Sub-rostrum. (/) Ditto, inside view of. 
(ff) Upper latus. 

Fig. 2. Pollicipes validus; all figures natural size. 

(a) Carina. (e) Scutum. 

(b) Ditto, inside view. (/) Ditto, inside view of. 

(c) Ditto, lateral internal view of. (ff) Ditto, inside view of another specimen. 

(d) Ditto, section of upper part. 

Fig. 3. Pollicipes gracilis. 

(a) Scutum, natural size. (b) Scutum, magnified, inside view. 

Fig. 4. Pollicipes dorsatus; all figures natural size, except (d), magnified twice. 

(a) Carina, inside view of. (d) Tergum, magnified twice. 

(b) Ditto, outside view of. (e) Tergum, natural size. 

(c) Ditto, section of upper part of. (/) Scutum, left-hand valve (vide supra). 

Fig. 5. Pollicipes striatus ; figures natural size, and magnified twice. 

(a) Carina. (b) Tergum; a left-hand valve (vide supra). (c) Scutum. 

Fig. 6. Pollicipes semiiatus ; magnified about ten times. 

Fig. 7. Pollicipes rigidus ; all figures thrice natural size, except (/), which is twice 

(a) Carina. (d) Scutum. 

(b) Ditto, side view of. (e) Tergum. 

(c) Ditto, section of lower part of. (/) Scutum, inside view. 

Fig. 8. Pollicipes fallax ; figures twice natural size. 

(a) Scutum. (b) Tergum. 

Fig. 9. Pollicipes elegans. 

(a) Carina, thrice natural size. (c) Scutum, twice natural size. 

(b) Ditto, section of. (d) Tergum, twice natural size. 

Fig. 10. Pollicipes Bronnii; figures magnified twice. 

(a) Carina. (d) Carina, section of, at one third of length from 

(b) Ditto, lateral view of. the apex. 

(c) Ditto, section of, near basis. 

Fig. 11. Pollicipes planulatus; Tergum, upper figure, natural size. 



Tab. W 




J.De CSffwerby fecit. 



TAB. V. 

Fig. 

1. Loricula pulchella ; natural size, as found embedded. 

2. Ditto Left-hand main valve, and scales of the peduncle, magnified 

three times. 

3. Ditto Right-hand main valve (scutum), and scales of the peduncle, 

magnified three times. 

4. Ditto Imaginary, restored figure. 



Tab. V. 




J. De (' . Sowerbv fecu 



A MONOGRAPH 



ON THE 



FOSSIL BALANID^E 



AND 



YERRUCID^E 



OF 



GREAT BRITAIN. 



BY 



CHARLES DARWIN, F.R.S., F.G.S. 



LONDON : 

PRINTED FOR THE PAL^EONTOGRAPHICAL SOCIETY. 

1854. 



J. E. ADLAKD, PRINTER, BARTHOLOMEW CLOSE. 



PREFACE. 

As the present short Monograph completes my work on British Fossil Cirripedes, 
I beg to be permitted again to have the satisfaction of returning my very sincere thanks to 
the many naturalists who have placed their collections at my disposal, and have given 
me the freest permission to use the specimens, in whatever manner I might find necessary. 
— My thanks are most especially due to Mr. Searles Wood, Mr. Bowerbank, and 
Sir Charles Lyell ; and to Mr. J. de C. Sowerby for the use of the original specimens 
figured in the ' Mineral Conchology.' I lie, also, under much obligation to Mr. D. Sharpe, 
Mr. Greenough, Mr. Smith of Jordan Hill, Professor Tennant, Mr. Charlsworth, 
Mr. F. Edwards, Dr. T. Wright, Professor Forbes, Professor Henslovv, M. Bosquet, and to 
many others. I must, also, be permitted to tender my grateful thanks to the Council of 
the Palseontographical Society for the very liberal manner in which they have allowed my 
two Monographs to be illustrated. 



INTRODUCTION. 



Cireipedia may be divided, as I have recently shown in a monograph on the Balanidse 
published by the Ray Society, into three Orders : of these, the Thoracica includes all 
ordinary Cirripedes, and all ever likely to be found fossil, and therefore the two other orders 
may be here passed over without notice. The Thoracica contains three Families : the 
Balanidae or Sessile Cirripedes, which in a recent state so abound on the shores of almost 
every quarter of the world, and which are so frequently found in Tertiary deposits ; the 
Verrucidae, which includes only a single genus very singular from its asymmetrical 
shell ; and the Lepadidae, or Pedunculated Cirripedes ; of the latter, the Fossil species have 
been already published by the Palaeontographical Society. The Balanidae and Verrucidae will 
be treated of in the following pages. As yet only sixteen species in these two families 
have been found fossil in Great Britain ; and of these sixteen, nine are still living forms. 
As the latter are known only imperfectly in their fossil condition, and as they have lately 
been described by me in full detail, I have thought it best here only to make a few remarks 
on such portions of the shells of each species as have hitherto been discovered, adding a 
few illustrations, such as appeared to me desirable. The extinct species will be fully 
described : of these, all the figures given are from British specimens. But of the species 
found both living and fossil, I have in several instances (always so stated) given drawings 
from recent specimens ; some of the valves either not having been found fossil, or 
found only in an imperfect and not characteristic condition. As so few species in the 
several genera are known in a fossil condition, I have thought it quite superfluous to give 
long generic descriptions, which would have required constant references to many species 
exclusively found living. 

In my former monograph on the Fossil Lepadidae, I remarked how much the natural 

1 



2 INTRODUCTION. 

history of Cirripedes has been neglected ; and this remark is eminently applicable to the 
Balanidae, or Sessile Cirripedes. Even the British recent species have not been well made 
out, and as for the fossil species, scarcely anything has been done, besides the publication of 
some figures, in very few instances accompanied by the details which are absolutely neces- 
sary for the identification of the species. 

Owing to the great variation in external characters, to which almost all the species are 
subject, and likewise in the case of the principal genus, Balanus, to its being a very natural 
genus, that is, to the species following each other in close order, it is not easy to 
exaggerate the difficulty of identifying the species, except by a deliberate examination of 
the internal and external structure of each individual specimen. Every one who has 
collected Sessile Cirripedes must have perceived to what an extent their shape depends on 
their position and grouping. The surface of attachment has a great effect on that of the 
shell ; for as the walls are added to at their bases, every portion has at one time been in 
close contact with the supporting surface ; thus I have seen a strongly-ribbed species 
(B. porcatus) and a nearly smooth species (B. crenatus) closely resembling each other and 
both having a peculiar appearance, owing to their having been attached to a pecten. 
Dr. Gray has pointed out to me specimens of the recent B. patellaris, curiously pitted like 
the wood to which they had adhered ; and numberless other instances might be added. 
Quite independently of the effect produced by the surface of attachment, the degree to 
which the longitudinal folds and ribs are developed on the parietes, is variable in most of 
the species, as in B. tintinnabulum and even in B. porcatus; the presence or entire 
absence of these ribs often surprisingly alters the whole aspect of the shell. The persistence 
of the so-called epidermis is in some degree variable, though this is of little importance in 
regard to fossil specimens. Again, some species in certain localities are all subject to the 
disintegration of the entire outer lamina of the walls ; and in such cases (as with 
B. p erf or at us) there is not the smallest resemblance between the corroded and perfect 
specimens. The size of the orifice, and consequently of the operculum, compared with the 
shell, varies accordingly as the shell is more or less conical or cylindrical ; in the 
latter case, the summits of the radii are generally more oblique and the orifice consequently 
more deeply toothed than in the more conical varieties. Size is a serviceable character in some 
cases, but very many specimens are required to ascertain the average or maximum size 
of each species, for there is no method of distinguishing a half-grown from a full-grown 
specimen ; and I believe, as long as the individual lives, so long does it go on growing. 
Colour is of very considerable service; but the majority of the species have their 
white or nearly white varieties, the latter being sometimes as numerous as the coloured 
ones. 

Besides the slight variation in the obliquity of the summits of the radii and alae, de- 
pendent on the more or less cylindrical form of the shell, in some species, as in B. 
tintinnabulum and porcatus, their obliquity also varies occasionally from unknown causes, 



INTRODUCTION. 3 

and thus greatly affects the general appearance of the shell. In some few species, as in 
B. perforatus, the radii are often either not at all developed, or are of very variable width ; 
in others, when the shell has become cylindrical, or when very old, the radii cease to grow, 
and from the disintegration of the whole upper part of the shell, with the continued growth of 
the lower part, the radii at last come to exist as mere fissures : I have seen instances of this 
in B. tintinnabulum and porcatus. Nevertheless, the obliquity of the upper margin, and 
the breadth of the radii are useful characters ; and still more useful is the fact whether the 
upper margins are smooth and arched, or straight and jagged. The fact of the terga being 
more or less beaked is useful : as is, likewise, the presence of striae, or furrows, or rows 
of pits, radiating from the apices of the scuta; but to ascertain the presence of these marks, 
it is almost invariably necessary to clean and examine the scuta with a lens ; these ridges 
and furrows, moreover, in some species, as is strikingly the case with B. tintinnabulutn, 
and in less degree with B. concavus, appear and disappear, and vary without any apparent 
cause. 

Now if we reflect that form, size, state and nature of the surface, presence of epidermis, 
relative size of the orifice, presence of longitudinal ribs, tint, and often the existence of any 
colour, are all highly variable in most of the species ; and that the obliquity of the summits 
of the radii, and the presence of longitudinal striae on the scuta, are variable in some of the 
fossil species, we shall perceive how difficult it must ever be to distinguish the species from 
external characters. Let no one attempt to identify the species of this genus, without being 
prepared to separate, clean, and carefully examine with a microscope the basis and parietes, 
and both the under and upper surfaces of the opercular valves ; for I feel convinced, that 
he will otherwise throw away much labour. Moreover, in many cases, it is almost neces- 
sary, on account of the variability of the characters, to possess several specimens. From 
these facts, I have not hesitated to use characters, for the identification of the species, which 
require close examination, though I would gladly have seized on external characters could 
I have found such even moderately constant. 

The least varying, and therefore most important characters, must be taken from the 
internal structure of the parietes, radii, and basis : not that these characters are absolutely 
invariable ; thus the porosity of the parietes is slightly variable in the recent B. glandula, 
and highly variable in the fossil B. unguiformis. The porosity of the basis is in some 
degree variable in B. spongicoia. Characters derived from the general shape, and from 
the ridges and depressions on the under side of the scuta and terga, especially of the 
scuta, are highly serviceable ; though even these are variable. The cause of the opercular 
valves offering more useful characters, as far as outline is concerned, than do the walls of 
the shell, is no doubt due to their being almost independent of any influence from the 
nature of the surface of attachment. Even the ridges and depressions on the under side of 
the scuta, which are in direct connexion with the muscles and soft parts of the animal, 
vary to a certain extent : thus the length and prominence of the adductor ridge is decidedly 



4 INTRODUCTION. 

variable in the fossil B. concavus and tintinnabulum / the size and form of the little cavity 
for the lateral depressor muscle varies in many species ; so does the exact shape and degree 
of prominence of the articular ridge. There is one character in the terga, which at first 
would be thought very useful, namely, whether an open longitudinal furrow, or a closed 
fissure runs down outside the valve from the apex to the spur ; but it is found that the furrow 
almost always gradually closes up during growth ; and as a consequence of this, the width 
of the spur compared to that of the whole valve, as well as its distance from the basi-scutal 
angle, and the form of its basal extremity, all vary in some degree. The length of the 
spur sometimes varies considerably, as in B. concavus. The summits of the radii are apt 
to be oblique in the young of some species, whereas they are generally quite square in the 
old of the same. In some species the scuta become longitudinally striated only with age ; 
on the other hand, in very young specimens of B. tintinnabulum, the scuta sometimes are 
deeply impressed by little pits placed in rows. I have already alluded to the longitudinal 
furrow on the tergum so entirely changing its character, owing to the edges becoming, 
during growth, folded inwards, and to the consequences which result from this. The inner 
lamina of the parietes generally loses, to a certain extent, its longitudinally ribbed character 
in old age. The basis is solid instead of being porose, in very young specimens of some 
species. In all the species, the carino-lateral compartments, in early age, are very 
narrow in proportion to the width of the lateral compartments ; and in all, at this early 
period, the operculum is large in proportion to the whole shell. 

Finally, I must state my deliberate conviction that Sessile Cirripedes can very seldom 
be satisfactorily identified in a fossil condition, without an examination of the opercular 
valves : hence when these have not been discovered, I have resolved, with some rare 
exceptions, not to attach a specific name to a shell without its operculum ; for thus, I 
believe, I should add to the number of useless synonyms, which, as we shall immediately 
see, already exist. Nothing, indeed, could have been easier than to have affixed names to 
many groups of specimens, having different aspects, but to feel sure that these were really 
distinct species requires better evidence than can be afforded by the shell, without the 
opercular valves. When the specimens are much fossilised, it is, indeed, difficult to make 
out the primary points of structure in the genus Balanus — namely, whether the parietes, 
radii, and basis are porose : to do this it is sometimes necessary to rub down, polish, and 
carefully examine, a transverse section of a piece of the shell. 

The ancient history of the Balanidse is a brief one. No Secondary species has hitherto 
been discovered ; in my former monograph on the fossil Lepadidse 1 I have shown that 
the negative evidence in this case is of considerable value, and consequently that there is 
much reason to doubt whether any member of the family did exist before the eocene period. 

1 Since the note to page 5 of that work was written, I have been informed that the so-called cretaceous 
Tubicinella maxima is not a Cirripede. 



INTRODUCTION. 5 

The existence of a Cretaceous Verruca is an apparent exception to the rule, as this genus has 
hitherto always been ranked among Sessile Cirripedes ; but Verruca, as we now know, must 
be placed in a family by itself, namely, the Verrucidee, quite distinct from the Balanidse. 
Balanus is the oldest genus as yet known ; it first appeared in Europe and North America, 
during the deposition of the eocene beds ; and was at that time, as far as our information at 
present serves, represented by very few species. In South America, one species of Balanus 
abounds in individuals in the ancient Patagonian tertiary formation. I have seen, in the 
British Museum, specimens said to have come from the eocene nummulitic beds, near the 
mouth of the Indus, belonging to that section of the genus, which has the walls and 
basis permeated by pores. Generally, the extinct forms belong to the section, which has 
the parietes not permeated by pores. During the miocene and pliocene ages, Sessile 
Cirripedes abounded. No extinct genus in this family has hitherto been discovered. It is 
singular, that though the Chthamalinae approach much more closely than do the Balaninse 
to the ancient Lepadidse, of which so many species have been found fossil even in the 
older Secondary formations, yet that only one species of one genus of this sub-family has 
been hitherto found in any deposit; and that species is the still existing Pachylasma 
giganteum, in the modern beds of Sicily. During the epoch of the Glacial deposits in 
Scandinavia, Scotland, and Canada, the still existing species seem to have abounded; 
and they attained larger average dimensions than the same species now do on the shores 
of Great Britain, or even on the shores of the northern United States, where the average 
size seems larger than on this side of the Atlantic. 

I already have given my reasons for very seldom naming any Sessile Cirripede 
without examining the opercular valves : it has been owing to this, as it appears to me, 
proper want of caution, that there are so many nominal species. Thus it is made to 
appear in catalogues, that the tertiary seas abounded with species of Balanus to an 
extent now quite unparalleled in any quarter of the world. Bronn, 1 for instance, in his 
invaluable ' Index Palaeontologicus,' gives the names of thirty-five Balani, found fossil in 
Europe, and I have not counted those found only in alluvial deposits, as they would 
certainly be the same as the still living species. Now I know only eleven recent Balani 
on the shores of all Europe, from the North Pole to lat. 30°; and of these I doubt whether 
B. balanoides and improvisus have been found fossil. In the Red Crag there is one extinct 
Balanus : in the Coralline Crag, which seems to have been very favorable to the existence 
of Cirripedes, there are six species of Balani, of which two are absolutely extinct, and 
one does not occur in any neighbouring sea : in the Eocene formations the species seem 
to have been rare, and I have seen only one, and that is an extinct form. Taking these 
several facts into consideration, and bearing in mind that Cirripedes usually range widely, 

1 To save any other person interested in fossil Cirripedia, going through the several works quoted by 
Bronn, I have given some remarks on his list of species, in an appendix at the end of the Balanidee, in my 
volume published by the Kay Society. 



6 INTRODUCTION. 

I do not believe, if all the specimens of Balani hitherto found in the several tertiary 
formations, from the Eocene to the Glacial deposits, throughout Europe, were collected 
together, they would amount to twenty species. I have myself seen, in a recognisable 
state, only twelve fossil species, of which five are extinct or not found in any neighbouring 
sea : I think it probable that three other recent species, viz., B. tulipi/ormis, perforatus, 
and amphitrite, may occur in the Mediterranean formations -. and this would make fifteen 
species. Therefore, in the above estimate of twenty species, five are allowed for species 
existing in European collections, but not hitherto seen by me ; and this, I believe, is a very 
full allowance. Consequently, even on the supposition that the five species just admitted 
as possibly existing in cabinets, and that the other five extinct species, which I have seen 
and examined, have all been previously named by other authors, a supposition excessively 
improbable, even then there would be fifteen superfluous names in Bronn. 

The following short table shows how the Balanidse and Verrucidse were represented in 
Great Britain, throughout the several Tertiary Stages. It includes all the sixteen species 
described in the following pages, with the exception of one, the cretaceous Verruca prisca, 
which is the only member of either family hitherto found in any Secondary deposit. 



Name. 


Living species, 

but found fossil 

in some tertiary 

deposits. 


Mammaliferous 

crag, and glacial 

deposits. 


60 
S3 
U 
CJ 

CO 


6J0 

CS 

u 

CJ 
CJ 

"ca 
o 
O 


cj 
p 

CJ 

CJ 

o 

m 


Balanus tintinnabulum 
calceolus 
spongicola 
concavus 
porcatus . 
crenatus . 
Hameri . 
bisulcatus 
dolosus . 
inclusus . 
unguiformis 

Aeasta undulata . 

Pyrgoma anglicum 

Coronula barbara . 

Verruca Stromia . 






; 

i 
i 

: 


: 

f 

i 


* 
* 

* 


* 

* 
* 

* 

* 

* 

* 


* 

* 
* 

* 

* 

* 

* 

* 

* 


a 


• 


Total 15, recent and extinct, 
found fossil in Great Britain, 
in some tertiary deposit 


9 


5 


8 


9 


1 



INTRODUCTION. 7 

As affording some standard of comparison by which to compare the number of species 
found fossil in any Tertiary deposit, in relation to the number of species probably existing 
in the neighbouring seas during the same epoch, I may state that there are now living and 
propagating "on the shores of Great Britain, eleven species belonging to the two Families 
included in the above table. In the Coralline crag, which seems to have been emi- 
nently favorable for the existence and subsequent preservation of Cirripedes, and which 
has been so well worked, nine fossil species of these two families, as may be seen in the 
table, have been discovered. 



NOMENCLATURE OF THE SHELL OF A SESSILE CIRRIPEDE. 



ARCHETYPE SHELL. Fig. 1. 

Orifice of shell, surrounded by the sheath. Sheath formed by the aim (a — a) and by portions of the 
upper and inner surfaces of the parietes (p — p). 



p p Parietes 



Basis 




n. b. In Balanus, and all known fossil genera, the Rostrum and Rostro-lateral compartments are confluent, 
and hence the Rostrum has the structure of Fig. 2. 



Fig. 2. 



COMPARTMENTS. 
Fig. 3. 



Fig. 4. 






Fig. 2. Compartment with two radii, serving, in fossil specimens, always as a Rostrum. 
Fig. 3 serves as a Lateral and Carino-lateral Compartment. Fig. 4 serves as a Carina. 



OPERCULAR VALVES. 



Fig. 5. Scutum (internal view). 

i V_ Tergal Margin 



Occi.udent Margin 

Cavity for Adductor 
Muscle 
Adductor Ridge 




'.!._ Articular Ridge 
■| Articular Furrow 

}... Cavity for the Lat. 
Depressor Muscle. 



Apex 
Articular Ridge ....J 
Articular Furrow ...J 
Scutal Margin 



Fig. 6. Tergum (internal view). 

Carinal Margin 




Crests for Depressor 

Muscle. 
Basal Margin 

Spur. 



Fig. 7. Tergum (external view). 
Beaked Apex 

-Carinal Margin 

Scutal Margin 



Spur 




Basal Margin 



INTRODUCTION. 



On the Names given to the different parts of Cirripedes. 

In my former volume I stated that I had found it indispensable, in part owing to 
the extreme confusion of the nomenclature previously used, to attach new names to several 
of the external parts of Cirripedes. Almost all these names are applicable to the Balanidse, 
or Sessile Cirripedes, and to the Verrucidse ; but a few additional names are requisite, 
which, together with the old names, will, I hope, be rendered clear by the accompanying 
woodcuts. In Sessile Cirripedes, the whole of that which is externally visible, may, for 
convenience sake, be divided into the operculum or opercular valves {valva operculares)> 
and the shell (testa), though these parts homologically present no real difference. The 
operculum is seated generally some little way down within the orifice of the shell ; but in 
very young specimens, and in Verruca, the operculum is attached to the summit of the 
shell, and in these cases the shell, without the operculum be removed, can hardly be said 
to have any orifice ; though, of course, the opercular valves themselves have an aperture 
for the protrusion of the cirri. 

The shell consists of the basis (called the support by some authors), and of the com- 
partments {testa valva), which in recent specimens vary from eight to four in number, and 
occasionally are all calcined together. 

The compartment, at that end of the shell (fig. 1) where the cirri are exserted through the 
aperture or lips of the operculum, is called the carina (fig. 4) ; the compartment opposite to 
it, is the rostrum (in all fossil specimens, like fig. 2), — these two lying at the ends of the 
longitudinal axis of the shell. Those on the sides are the lateral compartments ; that 
nearest the carina, being the carino-lateral (fig. 3) (testa valva carino-lateralis) , that 
nearest the rostrum, the rostro-lateral, and middle one simply the lateral compartment 
(fig. 3) : but these three compartments are rarely present together. The rostro-lateral com- 
partment, which always resembles fig. 2, and may be always known by having radii on 
both sides, is not known to occur in any fossil species ; and hence we are here only con- 
cerned with the lateral and carino-lateral compartments. The compartments are separated 
from each other by sutures, which are often so fine and close as to be distinguished with 
difficulty. The edge of a compartment, which can only be seen when disarticulated from 
its neighbour, I have called the sutural edge (acies suturalis). 

Each separate compartment consists of a wall (paries), or parietal portion (pp in figs. 1 
and 4), which always grows downwards, and forms the basal margin ; and is furnished on 
the two sides either with ala (fig. 4), or with radii (fig. 2), or with an ala on one side and 
a radius (fig. 3) on the other. 

The radius (adopting the name used by Bruguiere, Lamarck, and others) differs 
remarkably in appearance (though not in essence) from the wall or parietal portion, owing 
to the direction of the lines of growth and the state of its usually depressed surface. In 
the upper part, the radii overlie the alee of the adjoining compartments : in outline 

2 



10 INTRODUCTION. 

(r, fig. 2, 3), they are wedge-formed, with their points downwards ; their summits (and 
this is often a useful specific character) are either parallel to the basis, or as in fig. 1, 2, 
oblique. The radii are sometimes not developed. 1 

The alee (so called by Dr. Gray) are overlapped by the radii, and by part of the walls ; 
they usually extend only about half way down the compartment {a, fig. 3, and 4) ; their 
summits are either parallel to the basis or oblique. The alae of the several compartments, 
together with the internal, upper, thickened surfaces of the walls, against a shoulder of 
which the sutural edges of the alse abut, have been called (by Dr. Gray) the sheath {vagina). 
The upper and greater portion of the sheath is marked by transverse lines, caused by the 
exuviation of the opercular membrane, as that membrane may be called, which unites the 
operculum all round to the sheath, or upper internal surface of the shell. 

The carina has always two ales, as in fig. 4. 

The carino-lateral and lateral compartments have always an ala on one (the rostral) 
side, and a radius on the other (the carinal) side, as in fig. 3. 

The rostro-lateral compartment (not at present known to occur in any fossil) has always 
radii on both sides, as in fig. 2. 

The rostrum has normally ala on both sides, as in fig. 4 ; but in many recent, and 
all the fossil species yet known, it has radii on both sides, as in fig. 2, owing to its fusion 
with the rostro-lateral compartments on both sides. 

The walls of the shell, the basis, and the radii, are in very 
many cases composed of an outer and inner lamina, united 
together by septa ; a set of pores or tubes being thus formed. 

Basai edge of waii of compartment The points of the septa generally project beyond the lamina?, 

in Balamis tintirmabulum. a, a, outer * l o j i j ^ 

riS&^M^Mitto^'mnMMd an( l ave denticulated on both sides, as shown in the accompany- 

outer lamina;, with their ends denti- . . 

culated. lllg WOOOCllt. 

Operculum, or opercular valves. — These consist of a pair of scuta and a pair of terga. 
They are joined to the sheath of the shell by the opercular membrane. 

Scutum (fig. 5) : this valve is generally sub-triangular, and its three margins are 
the basal, the tergal, so called from being articulated with the tergum, and the occludent, 
so called from opening and shutting against the opposed valve. The angles are named, 
from the adjoining margins, as bad-tergal, &c. ; the upper angle being the apex. The 
scutum is ordinarily articulated to the tergum by an articular ridge [crista articularis) , 
running up to the apex of the valve, and by an articular furrow, which latter receives the 

1 The radii have been called by Ranzani and De Blainville "arese depressse" (the parietal portions of 
the compartments being the "arese prominentes"); by Poli, "areee interjectae;" by Gray, "sutures;" 
by Coldstream, "compartments of the second order," (the parietal portions being those of the first order) ; 
by some authors, "intersticia." I may here add that the scuta are the "ventral valves" of Gray, the 
"anterior" of Ranzani, and the "inferior opercular" of De Blainville: the terga are the "posterior valves" 
of Gray and Ranzani, but the " superior opercular" of De Blainville : the rostrum, on the other hand, is the 
" anterior valve" of Ferussac, and the "ventral" of De Blainville ; the carina being the " dorsal valve" of 
the latter author. 




INTRODUCTION. 1 1 

scutal margin of the tergum. The articular ridge, instead of projecting straight up from 
the valve, when the latter is laid flat on its external surface, often bends over to the tergal 
side, and is then said to be reflexed. On the internal surface of the valve there is almost 
always an adductor pit or cavity {fossa adductoris), for the attachment of the adductor 
scutorum muscle : this pit is often bounded on its tergal and basal sides, by a ridge, called 
the adductor ridge {crista adductoris), which, in its upper part, is often confluent with the 
articular ridge. Beneath the adductor ridge, in the basi-tergal corner of the valve, there 
is often a lateral-depressor pit (fossa muscidi lateralis depressoris), for the attachment of 
the so-called muscle ; and this pit is sometimes furnished with crests. 

Tergum, (figs. 6 and 7) : — this valve, also, has three margins, the scutal, basal, and 
carinal ; its upper end, or apex, is sometimes beaked ; on the basal margin a spur {calcar) 
depends ; the outer surface of the valve is depressed or longitudinally furrowed {sulcus 
longitudinalis) in the line of the spur. The part called the spur is often so broad, that the 
name becomes not very appropriate. The angles are denominated, from the adjoining 
margins, as basi-carinal, or basi-scutal angle, &c. On the under side, in the upper part, 
there is an articular ridge, and on its scutal side, an articular furrow, receiving the 
articular ridge of the scutum. In the basi-carinal corner of the valve there are often crests 
for the attachment of the tergal depressor muscle. 

Relative position of parts. — The centre of the generally flat basis, which is cemented 
to the supporting surface, is properly the anterior end, and the tips of the terga, often 
hidden within the shell, are properly the posterior end of the external covering; but I have 
found it more convenient to speak of the upper and basal surfaces and aspects, which 
hardly admit of any mistake. A line drawn from the centre of the basis, along the middle 
of the rostrum to the tips of the scula, shows the strictly medio-ventral surface of the shell ; 
and another line drawn from the centre of the basis, along the carina, to the tips of the 
terga, shows the strictly medio-dorsal line ; but from the crooked course of these lines, I 
have found it far more convenient to speak of the rostral and carinal end or aspect of the 
different parts of the shell. There has, moreover, been great confusion in these relative 
terms, as applied by different authors. 

When a sessile Cirripede is held in the position in which they have generally been 
figured, namely with the basis downwards and the scuta towards the beholder, then the 
right and left sides of the Cirripede correspond with those of the holder. 



Sub-class — Cirripedia. Order — Thoracica. 

Family — Balanid^e. 

Cirripedia sine pedunculo ; scuta et terga musculis depressoribus instructa ; reliance 
testa valva inter se immobiliter conjuncta. 

Cirripedia without a peduncle ; scuta and terga furnished with depressor muscles ; 
other valves united immoveably together. 

This family, which includes all true Sessile Cirripedes, may be divided into two very 
natural sub-families ; namely, the Balaninas and Chthamalinse ; but as not one member of 
the latter has been found fossil in Great Britain, and indeed only one, the Pachylasma 
giganteum, in any part of the world, viz., in the recent beds of Sicily, this sub-family of 
the Chthamalinse may be here passed over in silence. 



Sub-Family — Balanin^e. 

Rostrum cum radiis, sed sine alts : valves testa laterales omnes, ex uno latere alis, ex 
altero radiis instructa: parietes fere aut porosi, aut ad interior em superjiciem longitudi- 
naliter costati. 

Shell with the rostrum having radii, but without alas ; lateral compartments all having 
alse on one side and radii on the other side ; parietes generally either porose, or longi- 
tudinally ribbed on their inner surfaces. 

Genus — Balanus, Auct. 

Conopea (pars generis). Say. Journal Nat. Sc. Philadelphia, vol. ii, Part 2, 1822. 
Messdla (do.) Leach. Zoological Journal, vol. ii, 1825. 
Chirona (do.) J. E. Gray. Philosoph. Transacts., 1835, p. 37. 

Valva operculares infer se articulata, subtriangulares ; valva testa 6 ; basis calcarea aut 
membranacea. 



BALANUS. 13 

Scutum and tergum articulated together, sub- triangular ; compartments six ; basis 
calcareous or membranous. 

The genus Balanus already includes 45 species, recent and fossil, and consequently 
in my volume published by the Ray Society, I have divided the genus into sections, 
on characters derived from the porosity of the parietes, radii, and basis ; and on whether 
the basis be membranous or calcareous. But as here we have to describe or notice only 
eleven species, I have thought it more convenient to drop the sections, and in their place 
add a few words to each of the diagnostic characters. The genus is quite distinct from 
all the other genera of Sessile Cirripedes, with the exception of the sub-genus Acasta, 
from which its separation, it must be confessed, is in one sense artificial; for the 
species of this sub-genus graduate into those of Balanus (such as B. calceolus and its allies), 
which have their shells elongated in the rostro-carinal axis, and which live attached to 
Gorgoniae. These latter species have been generically separated by some authors from 
true Balanus ; but I have found it impossible to effect this ; and even the section of the 
genus, including these species, is hardly distinct enough from the adjoining sections. 
On the other hand, the sub-genus Acasta, in another sense, is a very natural one, inas- 
much as all its species are closely allied together in essential structure, in general 
appearance, and in habit ; and as the genus Balanus is already large, I have thought 
it best to adopt Acasta, which has been already admitted by many authors as a sub-genus. 
I need only further remark, that from reasons already assigned, I have thought it useless 
to give in this work long generic descriptions. 



1. Balanus tintinnabulum. Tab. 1, fig. la — Id. 

Lepas tintinnabulum. Linn. Syst. Naturae, 1/67. 

— — Ellis. Phil. Transact , vol. 50, 1758, Tab. 34, figs. 8 and 9. 

— — Chemnitz. Neues. Syst. Conch., 8 B. (1785), Tab. 97, 

figs. 828-831. 
Balanus tulipa. Bruguiere. Encyclop. Meth., 1789; sed non B. tulipa alba, in 
Chemnitz; nee non B. tulipa, O. F. Miiller, Zoolog. Dan. ; nee 
non B. tulipa, Poli, Test, ut Sicilise. 
— — G. B. Sowerby. Genera of Recent and Fossil Shells, Tab. Genus 

Balanus. 
Lepas ceispata (var.) SchrUer. Einleitung Conch., vol. iii, Tab. 9, fig. 21. 

— spinosa (var.) Gmelin. Linn. Syst. Nat. 

— tintinnabulum, spinosa, ceispata et poecata. W. Wood. General Con- 

chology, 1815, PI. 6, figs. 1, 2. PL 7, figs. 4, 5. PI. 8, figs. 1—5. 
Balanus tintinnabulum. Chenu. Illust. Conch. 

— d'Oebignii (var.) Chenu. Illust. Conch., Tab. 6, fig. 10, sed non Tab. 4, 

fig. 13. 

— ceassus. Sowerby (!) Min. Conch., 1818, Tab. 84. 



14 FOSSIL CIRRIPEDIA. 

B.parietibus etbasi et radiis poris perforatis : testa a rosed ad atro-purpur earn variante, 
soepe longitudinaliter virgatd et costata : orificio plerumque integro, inter dum dentato. Scuti 
crista articidari lata et rejlexd. Tergi margine basali plerumque in contrariis calcaris 
partibus rectam lineamformante. 

Walls, basis, and radii permeated by pores; shell varying from pink to blackish 
purple, often striped and ribbed longitudinally; orifice generally entire, sometimes toothed. 
Scutum with the articular ridge broad and reflexed. Tergum with the basal margin 
generally forming a straight line on opposite sides of the spur. 

Fossil in the Red Crag (Sutton). Mus. S. Wood, J. de C. Sowerby. Touraine (?) Mus. Lyell. 
Recent, on West Coast of Africa ; Madeira ; West Indies ; Cape of Good Hope ; Mouth of the Indus ; 
East Indian Archipelago ; Sydney, Australia ; Peru ; Galapagos Islands ; West Mexico ; California. 

Of this species I have seen several specimens, and fragments. Three of these are the 
original specimens figured in the Mineral Conchology, as B. crassus, an examination of 
which I owe to the great kindness of Mr. J. de C. Sowerby. Some specimens equally or 
more perfect are in Mr. S. Wood's collection. I have further seen a specimen from 
Touraine, which was presented to Sir C. Lyell by M. Dujardin, under the name of 
B.fasciatus, which I fully believe to be B. tinlinnabulum. None of these specimens had 
opercular valves, and therefore it is perhaps rash to assert quite positively that they are 
identical with B. tintinnabulum ; but, extraordinarily variable as this latter species is, yet, 
after having examined so many hundreds of recent specimens from all quarters of the 
globe, a sort of instinctive knowledge of general aspect is acquired, which makes me feel 
convinced that the fossils in question do really belong to this species. Moreover, the 
large shell, with its trigonal orifice passing into rhomboidal, — the smooth, broad, finely 
porose radii, with their summits not oblique, — the rather large parietal pores, — 
and the cancellated basis, are characters which hardly concur in any other species ; and 
those with w r hich these fossils might be confounded, are inhabitants of distant quarters of the 
world. Most of the recent varieties of B. tintinnabulum, and all the fossil specimens from 
the Crag, can be at once discriminated from B. tulipiformis (with which, at least in the 
Mediterranean deposits, it is likely to be confounded) by the summits of the radii extending 
from tip to tip of the adjoining compartments, and therefore not being oblique, as is 
always the case with the radii of B. tulipiformis. The largest fossil specimen which I have 
seen is nearly two inches in basal diameter, and nearly the same in height, and therefore 
about two thirds of the size of the largest living specimens. 

I have had engraved, from recent specimens, an internal view of the scutum and tergum, 
as these are likely hereafter to be found by searchers in the Crag deposits ; and I may 
refer to my Monograph on the Balanidae for their full description. It may be observed in 
the habitats given of the living specimens, that Madeira is the nearest point where the 
species now lives and propagates ; but specimens in full vigour are often brought to the 
British shores, attached to the bottoms of vessels. 



BALANUS. 15 



2. Balanus calceolus,* Tab. I, fig. 2a — 2d. 

Balanus calceolus keratophyto involutus (?) Ellis. Phil. Trans., vol.50 (1758), 

Tab. 34, fig. 19. 
Lepas calceolus (?) Pallas. Elench. Zooph., p. 198, (sine descript.) (1766). 
Conopea ovata (?) J. E. Gray. Annals of Philosophy, vol. 10, 1825. 

B. testes axe rostro-carinali elonyato ; basi cymbiformi ; parietibus et basi, sed non 
radiis, poris perf oralis. Scuto musculi depressoris lateralis fossa parvd, profunda. 

Shell with its rostro-carinal axis much elongated ; basis boat-shaped ; walls and basis 
porose, but not the radii. Scutum with the pit for the lateral depressor muscle small 
and deep. 

Fossil in Coralline Crag, attached to a Gorgonia ; Sutton ; Mus. S. Wood. 

Recent, attached to Gorgoniae, West Coast of Africa. Tubicoreen, near Madras. Mediterranean (?). 

I have seen only a single fossil specimen of this species, nearly half an inch in length, The 
shell was perfect, and a small portion of the Gorgonia yet remained attached to the grooved 
and boat-shaped basis. The opercular valves had been lost, but the shell in this instance 
is so peculiar, that it could only be confounded with the recent B. galeatus, cymbiformis, 
or navicula, and from all these it is easily distinguished by the parietes being permeated 
by pores. It is, of course, possible, that the opercular valves might present some new 
character, showing that this fossil, though agreeing with Bal. calceolus in its shell, yet 
was specifically distinct. I have given a drawing of the opercular valves from recent 
specimens, which have been fully described in my Monograph on the Balanidse. In 
regard to the shell, the fossil specimen could not be distinguished from the recent; and 
as it had to be broken, in order that its internal structure might be examined, I have 
thought it best to give a drawing from a perfect recent specimen. The spur of the 
tergum, in recent specimens, sometimes presents a singular character, in being irregularly 
toothed, and I have given a drawing (fig. 2d) of this variety, as it might perplex a collector. 

1 With respect to the nomenclature of this and three allied recent species, I must remark that in the 
published descriptions no allusion is made to any one of the characters by which alone they can be 
distinguished: hence I have been guided by geographical probabilities in assigning the specific name of 
calceolus to the present species, as Ellis's specimens came from the Mediterranean ; and that of galeatus to 
the North American and West Indian specimens, as Linnaeus' original specimens (according to a statement 
by Spengler) came from the West Indies. I have assigned new names to the two remaining East Indian 
species. 



16 FOSSIL CIRRIPEDIA. 



3. Balanus spongicola, Tab. I, fig. 3# — Se. 

Balanus spongicola. Brown's Illustrations of the Conchology of Great Britain (1827), 

pi. 7, fig. 6 : 2d edit. (1844), pi. 53, figs. 14—16. 

B. parietibus et basi, sed non radiis ports perforatis ; parietibus plerumque lavibus, 
roseis ; orifcio dentato ; scuto longitudinaliter striato ; tergum, apice producto, sine sulco 
longitudinali, calcare truncato, \ valvce latitudine, 

Parietes and basis, but not the radii, permeated by pores ; parietes generally smooth } 
shell pink ; orifice toothed ; scutum longitudinally striated ; tergum, with the apex pro- 
duced, without a longitudinal furrow ; spur truncated, about one third of width of valve. 

Fossil in Coralline Crag ; Sutton ; Mus. S. Wood. 

Recent on the South coast of England, and Tenby in South Wales ; Algiers ; Madeira ; Lagulhas Bank, 
Cape of Good Hope. 

I have seen only a single specimen of this species, which I picked out of a mass of 
specimens of the extinct Bal. inclusus, collected by Mr. Wood, in the Coralline Crag at 
Sutton. This one specimen was perfect, and included the opercular valves ; it even 
partially retained its rosy colour : it was "3 of an inch in basal diameter, and therefore 
exactly half the size of the largest recent specimen which I have seen. It was in every 
respect perfectly characterised. I have given drawings, external and internal, of the 
scutum and tergum from the fossil specimens. In the scutum, the adductor ridge is, 
perhaps, rather more prominent, and the pit for the lateral depressor muscle rather deeper 
than in recent specimens ; but these points are extremely variable. The tergum, in its 
outline, strictly agrees with the European recent specimens, and not with those varieties 
from the Cape of Good Hope and West Indies ; indeed, in the degree in which the basal 
margin on the carinal side of the spur slopes towards the spur, it even, perhaps, exceeds 
the European variety. These valves are fully described in my Monograph on the Balanidse. 
From the shell alone, as viewed externally, Bal. spongicola, even in its recent state, can 
hardly be distinguished from Bal. tulipiformis, or from some varieties of Bal. Capensis : 
I doubt whether this species could anyhow be distinguished in its fossil condition from the 
young of the fossil Bal. concavus, without the aid of the opercular valves. But in order 
to give an idea of its general appearance, and as I was compelled to disarticulate the com- 
partments of the one fossil shell, I have had a fine recent specimen from the Mediterranean 
engraved on an enlarged scale. 



BALANUS. 1 7 



4. Balanus concavus, Tab. I, fig. 4# — 4p. 

Balanus concavus. Bronn. Italiens Tertiar-Gebilde (1831) et Lethsea Geognostica, 
b. ii, s.1155 (1838), Tab. 36, fig. 12. 1 
— cylindkaceus, var. c. Lamarck. Animaux sans Vertebres (1818). 
Lepas tintinnabulum. Brocchi. Conchologia Sub-Appen., t. ii, p. 597, (1814). 

B. parietibus et basi, sed non radiis ports perforatis ; testa albo cum roseo aut obscure 
purpureo Jongitudinaliter pictd, interdum pure alba. Scuto longitudinaliter tenuiter striato : 
interne, adductoris crista admodum aut modice prominente. 

Parietes and basis, but not the radii, permeated by pores ; shell longitudinally striped 
with white and pink, or dull purple ; sometimes wholly white ; scutum finely striated lon- 
gitudinally ; internally, adductor ridge very or moderately prominent. 

Fossil in Coralline Crag, (Ramsholt and Sudbourne) rarely in the Red Crag (Sutton) ; Mus. 

5. Wood, Bowerbank, Lyell, J. de C. Sowerby, Tennant. Sub-Appenine formations, near Turin, Asti, and 
Colle in Tuscany, Mus. Greenough, &c. Tertiary bed, near Lisbon, Mus. D. Sharpe and Smith. Bordeaux (?) 
Mus. Lyell. Tertiary beds, Williamsburg ; and Evergreen, Virginia, Mus. Lyell. Maryland, Mus. Krantz. 
Pleiocene formations 2 near Callao, Peru, Mus. Darwin. 

Recent at Panama; Peru; S. Pedro, California; Philippine Arch.; Australia. 

This species has caused me much trouble. It will be convenient first to make a few 
remarks on the recent specimens ; I examined several from Panama and California, which, 
though differing greatly in colour, resembled each other in their scuta having the adductor 
ridge extremely prominent, and in having (Tab. I, fig. 4«) an almost tubular cavity for 
the attachment of the lateral depressor muscle, — characters which at first appeared of high 
specific value ; but I soon found other specimens from Panama in which these peculiarities 
were barely developed. I then examined a single specimen from the Philippine Archi- 
pelago, resembling in external appearance one of the Panama varieties, but differing in the 
scuta being externally strongly denticulated in lines instead of being merely striated, — in 
the adductor ridge being far less prominent, — and in the spur of the tergum being broader 
and more truncated ; I therefore considered this as a distinct species. I then examined a 
single white rugged specimen from the coast of Peru, which differed from the Philippine 
specimen in the shape of the well-defined denticulations on the scuta, and in some other 
trifling respects, and in the segments of the posterior cirri bearing a greater number of 
spines ; with considerable doubt, 1 also named this as distinct. But when I came to 

1 I suspect that B. pustularis, miser, and zonarius, all figured by Munster, in his 'Beitrage,' b. iii, 
Tab. 6, may be this species. 

2 I procured this specimen from the Island of S. Lorenzo, off Callao; it was imbedded, together with 
seventeen species of recent shells and with human remains, at the height of eighty-five feet. 



18 FOSSIL CIRRIPEDIA. 

examine a large series of fossil specimens from the Coralline Crag of England, and others from 
northern Italy, from Portugal, and from the southern United States, I at once discovered 
that the form of the denticuli on the striae of the scuta was quite a worthless character, — that 
in young specimens the scuta were simply striated, — that the prominence of the adductor 
scutorum riclge and the depth of the cavity for the lateral depressor muscle varied much 
(as in the case of the recent specimens), owing apparently to the varying thickness of the 
valve, — that in the terga the spur varied considerably in length and breadth, the latter 
character being in part determined by the varying extent to which the edges of the longi- 
tudinal furrow are folded in, — and lastly, that in young specimens the basal end of the 
spur is much more abruptly truncated than in the old. Hence I was led to throw the 
three recent forms, originally considered by me as specifically distinct, into one species ; 
but I may repeat that this considerable variation in the prominence of the adductor ridge, 
and in the depth of the pit for the lateral depressor muscle — the pit in some cases 
becoming even tubular — is a very unusual circumstance. 

With respect to the fossil specimens from the above stated distant localities, I consider 
them as belonging to one species, though they vary considerably in several points of 
structure. When compared with the recent specimens, they differ from them in often 
attaining a considerably larger size ; in the parietes being generally longitudinally ribbed, 
as in the case of the Coralline Crag specimen (Tab. I, fig. 4a), and in the radii often having 
more oblique summits. Some of the specimens from the United States, have strong 
rugged, depressed shells, frequently resembling, to a curious degree, Bal. porcatus. On 
the other hand, considering the many points of identity between the fossil and the recent 
specimens, I have concluded, without much doubt, that they ought all to be classed 
together. In the Coralline Crag specimens, the spur of the tergum (Tab. I, fig. 4jp) is 
unusually long and narrow; it is broader in the Italian specimens (4o), and either short 
(4/k) or long in the United States specimens. The scuta of the Lisbon specimens are 
remarkable for the great prominence of the adductor ridge, and for the depth of the lateral 
depressor cavity, as in most, but not in all, of the Panama specimens. The opercular valves, 
however, of some of the specimens from all these several distant localities are identical 
with the recent ones from the coast of America. I have entered into the above parti- 
culars, on account of, in the first place, its offering an excellent example how hopeless 
it is in most cases to make out the species of this difficult genus without a large series 
of specimens ; secondly, as showing how the characters alter with age ; and thirdly, as a 
good instance of the amount of variation which seems especially to occur in most of the 
species which have very extensive ranges. 

Some of the pink-striped Panama varieties, though having a somewhat different aspect, 
can be distinguished from certain varieties of B. amphitrite only by their scuta being lon- 
gitudinally striated, — a character in this species variable in degree, and in most cases of 
very little value. Some of the other recent varieties, however, are sufficiently distinct from 
B. amphitrite ; and the great fossil Coralline Crag specimens, which stand at the opposite 



BALANUS. 19 

end of the series of varieties, with their ribbed walls, very oblique radii, and coarsely 
striated scuta, are extremely unlike B. amphitrite. 

With respect to the nomenclature of the present species, I have little doubt that I 
have properly identified the Italian fossil specimens with B. concavus of Bronn, who has 
given a very good figure of this species in his ' Lethaea Geognostica ;' but it must 
be confessed that the longitudinal striae on the scuta are not there represented. Consi- 
dering the large size and frequency of this species in Europe and in the United States, it 
has probably received several other names besides the two synonyms, quoted at 
the head of this description. I should add that the true B. cylindraceus (not var. c) of 
Lamarck, according to the plate given by Chenu in his ' Illust. Conch.' is the B. psittacus 
of South America. I have seen in collections specimens of B. concavus labelled as 
B. tulipa of Poli (B. tulipiformis of my Monograph), — a very natural mistake, without 
the opercular valves be carefully examined. 

General Appearance. — Shell conical (fig. 4a), often steeply conical (fig. 4c), but 
sometimes depressed and smooth (fig. 4<d) ; orifice generally rather small, varying from 
rhomboidal to trigonal, with the radii narrow, and generally in the fossil specimens very 
oblique ; surface generally smooth, sometimes rugged, and in the Coralline Crag specimens 
commonly ribbed longitudinally, the ribs being narrow. In the recent specimens the colour 
is various, either dull reddish-purple with narrow nearly white, or wider dark longitudinal 
bands ; or, again, pale rosy-pink with broad white bands ; or lastly, wholly white. The 
radii are either darker or paler than the parietes. The opercular valves are either dark 
purple or nearly white. Pale pink and white stripes are visible on some of the Italian and 
Portuguese tertiary specimens ; and in most of the fossils the sheath is tinged dull red. 

Dimensions. — The largest actually recent specimen which I have seen, from the 
Philippine Archipelago, had a basal diameter of 1'2 of an inch; the Peruvian pleistocene 
specimen is 1*7 in diameter; specimens from the crag and from the Italian deposits, how. 
ever, sometimes slightly exceed two inches in basal diameter, and three in height. 

Scuta : these in young and moderately-sized specimens are striated longitudinally 
(fig. 4/), sometimes faintly, but generally plainly, causing the lines of growth to be beaded ; 
but in large and half-grown specimens, the lines of growth are often extremely prominent, 
and being intersected by the radiating stria?, are converted into little teeth or denticuli. As 
the stria? often run in pairs, the little teeth frequently stand in pairs, or broader teeth have 
a little notch on their summits, bearing a minute tuft of spines. In very old and large 
specimens, the prominent lines of growth are generally simply intersected by deep and narrow 
radiating striae (tab. I, fig. 4j»). In one case, a single zone of growth in one valve was quite 
smooth, whilst the zones above and below were denticulated. The valve varies in thickness, 
which I think influences the prominence of the lines of growth and the depth of the striae. 
These striae often affect the internal surface (fig. 4//) of the basal margin, making it bluntly 
toothed. The articular ridge (fig. 4n), is rather small, and moderately reflexed. The adductor 
ridge (as already stated) varies remarkably ; in most of the recent Panama specimens (fig. 4#), 



20 FOSSIL CIRRIPEDIA. 

and in the fossils from Portugal, it is extremely prominent, and extends down to near the basal 
margin ; in other specimens it is but slightly prominent, as in those from the Crag (4/) ; 
it is short, but rather prominent in the specimens (4//) from Maryland ; whereas it is very 
slightly prominent in the specimens from Virginia. The cavity for the lateral depressor, 
also, varies greatly ; it is often, as in the recent specimens, bounded on the side towards the 
occludent margin by a very slight straight ridge, which occasionally folds a little over, 
making almost a tube; this, at first, I thought an excellent specific character, but far from 
this being the case, the cavity often becomes, in recent specimens as well as in the crag 
specimens (4/), wide, quite open, and shallow. The whole valve in the Crag specimens 
(fig. 4e) is apt to be more elongated than in the recent or Portuguese specimens (fig. 4/), 
and especially than in the Maryland (fig. 4/t) specimens. 

Terga very slightly beaked ; the surface towards the carinal end of the valve, in 
some of the fossil specimens, is feebly striated longitudinally. There is either a slight 
depression (fig. 4k), or more commonly a deep longitudinal furrow (fig. 4g, 46) with 
the edges folded in and touching each other, extending down the valve to the spur, and 
causing the latter to vary in width relatively to its length. When the furrow is closed in, 
the spur is about one fourth of the entire width of the valve, and has its lower end 
obliquely rounded, and stands at about its own width from the basi-scutal angle : when 
there is only a slight depression and no furrow (as is always the case with young specimens, 
and in the specimens {41c) from Maryland), the spur is broader, equalling one third of the 
width of the valve, with its lower end almost truncated, and standing at about half its own 
width from the basi-scutal angle. But the absolute length of the spur, also, varies consider- 
ably in the Coralline Crag specimens ; it is often very long, (fig. 4g) compared to the whole 
valve. In many Italian specimens (4o) it is long and broad. The basal margin of the 
valve on the carinal side of the spur is sometimes slightly hollowed out ; and when the longi- 
tudinal furrow is closed, this side slopes considerably towards the spur. Internally, the 
articular ridge and the crests for the tergal depressor muscles are moderately prominent. 

Parietes, the longitudinal septa sometimes stand near each other, making the parietal 
pores small. The radii have oblique summits, but to a variable degree; their septa are 
unusually fine, and are denticulated on their lower sides; the interspaces are filled up 
solidly. The alee have their summits very oblique, with their sutural edges nearly or 
quite smooth. In most of the fossil specimens (Tab. I, fig. 4b, r), and slightly in some of the 
recent specimens, the surface of the sheath presents an unusual character, in a narrow, lon- 
gitudinal, slightly raised border, running along the sutures, on the rostral side of each suture. 

Basis thin, porose; sometimes with an underlaying cancellated layer. 

All the recent specimens which I have seen, were, with one exception, attached to 
various shells and crabs, and to each other. The tertiacy specimens are often congregated 
together into great masses. Including the recent and fossil specimens, this species encircles 
the globe. During the miocene period it seems to have been the commonest existing 
Sessile Cirripede ; now, it does not appear to be common, excepting, perhaps, at Panama. 



BALANUS. . 21 



5. Balanus porcatus. Tab. I, fig. ba — hg. 

Balanus porcatus. Emanuel da Costa. Hist. Nat. Test. Brit., p. 249, (1778). 
Lepas balanus. Linn. Syst. Naturae, (1767). 

— — Born. Testacea Mus. Cees. Desc., Tab. 1, fig. 4, (1780). 

Chemnitz. Syst. Conch., 8 Band., Tab. 97, fig. 820, (1785). 
Balanus arctica patelliformis. Ellis. Philosoph. Transact., vol. 50, Tab. 34, 

fig. 18, (1758). 

— sulcatus. Bruguiere. Encyclop. method., Tab. 164, fig. 1, (1789). 
Lepas costata and Balanus. Donovan. British Shells, 1802-1804, Tab. 30, fig. 1, 2. 
Lepas Scotica. W. Wood. General Conchology, PI. 6, fig. 3, sed non Lepas balanus, 

PL 7, fig. 3, (1815). 
Balanus angulosus. Lamarck (1818), in Chenu, Illust. Conch., Tab. 11, fig. 11. 

— tesselatus. Sowerby (!) Mineral Conchology, Tab. 84, (1818). 

— Scoticus. Brown. Illust. Conch. Great Britain, PI. 7, fig. 2, sed non 

PL 6, fig. 9 et 10 (1827) : 2d edit., PL 53, fig. 1-3, 22, 23 et 
PL 54, fig. 1-3. 

— geniculates. Conrad. Journal Acad. Philadelphia, vol. vi, part 2, p. 265 

(1830), Tab. 11, fig. 16. 

— — Aug. Gould (!) Report on the Invertebrata of Massachussetts, 

fig. 9 (1841). 

B. parietibus, sed non bast, ports per/oratis ; testa alba, pleritmque longitudinaliter 
acute costata; radio rum mar ginibus superioribus pame basi parallelis : scuto longitudinaliter 
striato ; tergi apice producto, purpureo. 

Parietes, but not the basis, permeated by pores ; shell white, generally sharply ribbed 
longitudinally ; radii with their summits almost parallel to the basis. Scutum longitudi- 
nally striated ; tergum with the apex produced and purple. 

Fossil in the Glacial deposits of Scotland (Isle of Bute), of Uddevalla, and (Beaufort) Canada. In the 
Mammaliferous Crag (Bramerton, Thorpe) and Red Crag (Sutton) ; Mus. Lyell, J.deC. Sowerby, S. Wood, 
Bowerbank, &c. 

Recent, England, Ireland, Scotland, Shetland Islands, Iceland, Davis's Straits, 66° 30' N. ; Lancaster 
Sound, 74° 48' N. Maine and Massachussetts, United States. China (?) In deep water, commonly 
adherent on shells, Crustacea, and rocks. 

This species can be at once distinguished from all the foregoing by the basis being 
solid or not perforated by pores ; and from all the following species, with the exception of 
B. crenatus, by the parietes having large square pores or tubes. From B. crenatus, this 
species can be distinguished by its longitudinally striated scuta, purple-beaked terga, and 
by the peculiar structure, immediately to be described, of its parietal pores ; and in most 
cases even by its general aspect, larger size, and ribbed walls. When, however, 
B. porcatus and crenatus have grown together on the same irregular surface, for instance, 
on a Pecten, they sometimes resemble each other in a very deceptive manner. The 
opercular valves have not certainly been found fossil, but I have given drawings from recent 
specimens. 



22 FOSSIL CIRRIPEDIA. 

The parietes, (the basal margin of a small portion is represented at Tab. I, fig. 53,) are per- 
forated by large square longitudinal tubes : in the upper part these are filled up solidly without 
transverse septa ; the longitudinal septa between the tubes are finely denticulated at their 
bases, and the denticuli extend unusually close to the outer lamina. In very young speci- 
mens the inner lamina of the parietes is ribbed, in lines corresponding with the longitudinal 
septa, as in the case of other species of the genus ; but in medium and large-sized 
specimens, there are between such ribs from one to four smaller ribs, which do not 
correspond with any longitudinal septa ; these are finely denticulated at their bases, and 
may be considered as the representatives of longitudinal septa which have not been 
developed and reached the outer lamina. I have seen no other instance of this structure, 
namely, the presence of a greater number of ribs on the inner (upper in fig. hb) lamina of 
the walls than there are longitudinal septa. As the parietes rest on the basis, the cir- 
cumference of the latter becomes marked in a very peculiar manner (fig. 5c), by the basal 
edges of the parietal septa. Hence the basis of this species can be distinguished from that 
of every other sessile cirripede : its circumference is plainly impressed by the main parietal 
septa which connect the inner and outer lamina? of the walls ; and between these marks 
there are two or three smaller impressions of the so-called representative septa, which do 
not extend beyond the impression of the basal edge of the inner lamina. The upper sur- 
face of the middle part of the basis (more especially when slightly disintegrated) is faintly 
striated in radiating lines, of which the stronger lines are prolonged from the circumferential 
marks left by the main parietal septa, and the weaker lines from the marks left by the 
representative septa. 

Dimensions. — The largest recent specimens which I have seen from great Britain or 
Ireland, have been 1*3 of an inch in basal diameter: in Mr. Cuming's collection, however, 
there is one much depressed specimen from the Shetland Islands, 2T in basal diameter; 
a regularly conical specimen from the coast of Massachusetts attains a nearly equal 
diameter. But out of the glacial deposits in the Isle of Bute, several specimens 
have this same diameter, namely, two inches, and are even more steeply conical, being 
1*85 in height; some glacial specimens from Uddevalla and Canada, in Sir C. Lyell's 
collection, are 1*7 in basal diameter. Hence it appears, as we shall presently see is 
likewise the case with B. crenatus and Hameri, that northern specimens, and those from 
the United States and from the Glacial deposits, often exceed in dimensions those now 
living on the coasts of Great Britain and Ireland, or those found in the Crag. 

This species is very common in the glacial deposits of Uddevalla, of Skien in Norway, 
and of Canada, and is associated with the same species, namely, B. crenatus and Hameri, 
as in the living state : I have seen, also, as just stated, specimens from the same formation 
in the Island of Bute, Scotland. I have examined numerous specimens from the Marn- 
maliferous Crag, and a few from the Red Crag of England. I owe to the kindness of 
Mr. J. de C. Sowerby an inspection of the original specimens of B. tesselatus of the 
Mineral Conchology, which is certainly the present species. 



BALANUS. 23 



6. Balanus crenatus. Tab. I, fig. 6a — 6g. 

B. crenatus. Bruguiere. Encyclop. Method, (des Vers) 1789. 

Lepas foliacea, var. a. Spengler. Skrifter af Naturhist. Selskabet, b. i, 1790. 

— borealis. Donovan. British Shells, PL 163, (1802-1804). 
B. rugosus. Pulteney (?) Catalogue of Shells of Dorsetshire, 1799. 

— Montagu (?) Test. Brit., 1803. 

— Gould (!) Report on Invertebrata of Massachusetts (1841), fig. 10. 
B. glacialis (?) J. E. Gray. Suppl. Parry's Voyage, 1819. 

B. elongatus (!) clavatus (!), auctorum variorum. 

B. parietibus, sed non basi pons perforatis ; testa alba ; radiorum marginibus superi- 
oribus obliquis, asperis, rectis ; scuto sine adductoris crista; tergi calcare rotundato. 

Parietes but not basis permeated by pores ; shell white ; radii with their oblique 
summits rough and straight; scutum without an adductor ridge; tergum with the spur 
rounded. 

Fossil in glacial deposits of Scandinavia and Canada, Mus. Lyell; in the mammaliferous and Red 
(Sutton) and Coralline Crags; Mus. S. Wood, J. de C. Sowerby, Bowerbank, &c. Miocene formation, 
Germany, Mus. Krantz. 

Recent in Great Britain, Scandinavia, Arctic Regions as far as Lancaster Sound, in 74° 48' N. ; 
Behring's Straits ; United States ; Mediterranean ; West Indies ; Cape of Good Hope. Generally attached 
to shells and Crustacea in deep water. 

Under the last species I have shown that the porose parietes, but solid basis, distin- 
guish this species easily from all the others, with the exception of B. porcatus, from 
which it can readily be known by the characters of its opercular valves, as already there- 
under stated. Judging by external appearances alone, which ought never to be trusted to 
in the identification of any sessile cirripede, this species might easily be confounded with 
Bat. dolosus, found fossil in the same deposits. 

This species presents a great diversity of external aspect : I have had figured (Tab. I, 
fig. 6a) one of the commonest appearances presented by it ; but frequently the shell is 
quite smooth and depressed, or extremely much elongated and cylindrical, or even club- 
shaped, The basis is generally thin and slightly furrowed in lines radiating from the 
centre, but it is not permeated by pores ; when, however, in large and old specimens it 
becomes thicker, as in Tab. I, fig. 6c, its edge is very distinctly pitted by little hollows, 
which might sometimes be easily mistaken for the orifices of pores : the absence of pores 
is a very important character in the diagnosis of B. crenatus. The basis is less firmly 
attached to the supporting surface than is usual with most cirripedes, and consequently it 
often separates from it together with the parietes. With regard to the opercular valves 
{6d — 6g) drawn from recent specimens, I need here only state that the most conspicuous 



24 FOSSIL CIRRIPEDIA. 

characters are the large articular ridge to the scutum, and the reflexed apices of all four 
valves, though this latter character is highly variable. I must refer to my Monograph on 
the Balanidse for a full description of these valves. 

The largest recent British specimen which I have seen was only "55 of an inch in basal 
diameter : specimens from Greenland and the northern United States, frequently attain a 
diameter of three-quarters of an inch, and I have seen one single sowewhat distorted 
specimen actually 1*6 of an inch in basal diameter. Where individuals have grown 
crowded together, their length is often twice, and even occasionally thrice as great as their 
diameter; thus I have seen a recent Greenland specimen l - 6 of an inch in length, and 
only *75 in diameter. This species, in its recent state, as may be seen under the 
habitats, has an enormous range. I have felt myself unwillingly compelled to admit that 
it ranges from the Arctic Regions in 74° 48' N. to the Mediterranean, the West 
Indies, and Cape of Good Hope. That this species should live in the tropical seas is the 
more surprising, as the large size of the specimens in the northern seas and in the glacial 
deposits, might fairly have been supposed to have indicated special adaptation for a cold 
climate. This great geographical range, however, of the species accords with its range in 
time from the present day to the Coralline Crag period. The specimens from the glacial 
deposits which I have examined, chiefly in Sir C. Lyell's collection, are very fine and large, 
and appear, on an average, to attain as large or larger dimensions than the recent specimens 
from the United States ; they are often associated, like the now living individuals, with 
B. porcatus and Hameri : they come from the well-known formation of Uddevalla, and 
from Canada. There are well-characterised specimens in the mammaliferous Crag, at 
Brain erton and near Norwich, in Sir C. Lyell's and Mr. Wood's collections, and from 
Sutton and other places in the Red Crag of the eastern shores of England : these speci- 
mens are not only smaller than the glacial, but than the recent English specimens ; 
for the largest Crag specimens which I have seen had a basal diameter *5 of an inch, '3 to '4 
being their ordinary size. The specimens which I have seen from the Coralline Crag, and 
some others sent me by Krantz from the miocene formation of Flonheim bei Abzei, in 
Germany, had not their opercular valves, yet I cannot doubt, considering how few species 
there are having porose walls and a solid basis, that I have rightly identified these specimens 
as belonging to B. crenatus. 



7. Balanus Hameri, Tab. I, fig. la— Id., Tab. II, fig. la, lb. 

Lepas Hameb.1. Aseanius. Icones rerum naturalium, Tab. 10, 1767. 

— tulipa. O. F. Midler. Prodromus. Zoolog. Dan. 1776 ; sed non L. tulipa, in 

Poli, Test, ut Sicilise ; necnon B. tulipa, in Bruguiere, Encyclop. 
method. ; necnon B. tulipa, in Sowerby, Genera of Shells. 

— tulipa alba. Chemnitz. Syst. Conch., Tab. 98, fig. 832. 

— foliacea. Spengler. Skrivter af Naturhist. Selskabet, 1 B. 1790. 



BALANUS. 25 

Balanus candidxjs. (Tab. emendata) Brown. Conch. Great Britain (1827), Tab. 6, 

figs. 9 and 10, and 2d edit. Tab. 54, figs. 9-12. 
— tulipa. Lyell. 1 In Phil. Transact., 1835, p. 37, Tab. 2, figs. 34-39. 

B. nee parietibus, nee basi, nee radiis ports perf oralis ; testa alba; radiorum margini- 
bus superioribus obliqicis, Icevibus, arcuatis ; aciebus suturalibus lavibus ; scuto angusto 
longitudinallter , debiliter striato ; tergi calcare angusto, rotundato. 

Parietes, and basis, and radii not permeated by pores ; shell white ; radii with their 
oblique summits smooth and arched; sutural edges smooth; scutum narrow, feebly 
striated longitudinally ; tergum with the spur narrow, rounded. 

Fossil in Red Crag (Sutton), Mus. S.Wood Doubtfully in the Glacial beds of Scotland. In the 
Glacial deposits at Uddevalla, in Sweden; and Beaufort, Canada, Mus. Lyell. Banks of the Dwina, 
Russia, Mus. Murchison. Greenland, "in blue clay," according to Spengler. 

Recent on the Coast of Yorkshire ; Scotland ; Galway, Ireland ; Isle of Man, and Anglesey, twelve 
fathoms. Generally in deep water ; not very common. George's Bank, Massachussetts, United States. 
Iceland, Finmark, and the Faroe Island, according to Spengler. Attached to Crustacea, mollusca, stems of 
fuci, and stones ; often associated with B. porcatus and crenatus. 

I have seen, in Mr. Wood's collection, from the Red Crag, fine and perfectly preserved 
specimens of a rostrum, and of a lateral compartment. The latter was three inches in height, 
and, including the alae, one inch in width. I have also seen a specimen said doubtfully to 
have come from the glacial beds of Scotland. As it is so very common in the deposits of 
this same age in other countries, no doubt it will hereafter be found more plentifully in 
Scotland, and probably in the mammaliferous Crag of England. Balanus Hameri is a 
very fine species ; I have seen a recent specimen from the coast of Yorkshire, two inches 
in diameter, and one inch and three-quarters in height : another specimen was three inches 
in height. The specimens in the glacial deposits, seem to have acquired larger dimen- 
sions : a compartment from Uddevalla being nearly four inches in height. The white 
colour, smooth surface, and regularly arched radii, give to the shell a very elegant appear- 
ance, which has appropriately been compared to that of a white tulip. The diagnosis of 
this species is easy ; the walls as well as the basis being solid or not porose, serve to dis- 
tinguish it from all other forms except certain varieties of Bal. ungiciformis, and by several 
minor characters, such as the finely striated and more elongated scuta, &c, Bal. Hameri, 
can be discriminated from B. unguiformis. 

1 Sir C. Lyell remarks that this is apparently the B. Uddevallensis, (Linn.), of Swedish lists of fossils. 
Prof. E. Forbes has shown ('Mem. Geolog. Survey of England,' vol. i, p. 364) how this name arose, from 
a short description, prior to the introduction of the binomial system, " Lepas quae Balanus Uddevallensis," 
given by Linnaeus in his Wast-Gotha Resa, in 1747. 

For the reference to Ascanius' work, which is on the binomial system, and subsequent to the 10th 
edit, of Linnaeus in 1758, I am greatly indebted to Mr. Sylvanus Hanley. Had it not been for this 
gentleman, I should have used Miiller's name of B. tulipa as the first name. 

4 



26 FOSSIL CIRRIPEDIA. 

In the recent condition, the compartments of the dead shell fall apart with singular 
facility; and Sir Charles Lyell has remarked ('Philosophical Transactions,' 1835, p. 37) 
that in the glacial deposits of Scandinavia, the shell is never found whole, but the separated 
compartments in abundance : it appears, also, that the basis likewise easily separates from 
its support. The extreme edge of the basis is finely crenated, and not pitted as in Bal. 
crenatus ; the crenations or teeth are produced by the edge of the basis fitting in between 
the longitudinal septa on the internal surface of the parietes. There is one peculiarity in 
the alae of this species in its recent state, which I have observed in no other species, and 
which can be distinguished in some of the fossil specimens, as in Tab. II, fig. 13, — 
namely, the presence of an excessively fine linear furrow running along the sutural edge, 
a little towards the inner side, and filled (in the recent state) with a yellow ligamentous 
substance. 

In regard to the opercular valves, (Tab. I, fig. la — Id, drawn from recent and glacial 
specimens,) I need here only mention, that in the scuta, their flatness, elongation, and delicate 
longitudinal strise, are their chief characteristics. In very old and large specimens of the 
terga (as in the specimen, fig. Id, figured from Uddevalla), the basal margin on the carinal 
side of the spur slopes down towards it in a remarkable manner. 



8. Balanus bisulcatus, Tab. II, fig. 2a — 2/t. 

Balanus sulcatinus (?) Nyst, apud D'Omalius (sine descript. ant tabula), Geologie 

de Belgique, 1853. 1 

B. nee parietibus, nee radiis poris perf oralis j basi poris magnis perforata ; radiorum 
marginibus superioribus obliquis, lavibus ; aciebus suturalibus lavibus ; scuto angusto, sulci s 
longitudinalibus 2 ad 4 ; tergi calcare brevissimo dimidid valvce latitudine. 

Neither walls nor radii permeated by pores ; basis permeated by large pores ; radii 
with their upper margins oblique and smooth ; sutural edges smooth ; scutum narrow, 
with from two to four longitudinal furrows ; tergum with the spur very short, broad as 
half the valve. 

Var. plicatus (fig. 2c), with the walls deeply folded ; radii narrow, with their upper 
margins very oblique. 

Fossil in Coralline Crag ; Ramsholt, Gedgrave, Sutton ; Mus. S. "Wood, Bowerbank, J. de C. Sowerby. 
Rauville, dans )e Cotantin, Mus. G. B. Sowerby. Var. plicatus, Coralline Crag, Sutton, Mus. S. Wood, 
Bowerbank. Bolderberg, near Hasselt, Belgium, Mus. Bosquet. 

1 I am indebted to M. Bosquet for a specimen, bearing this name and reference, found in the 'Systeme 
Bolderien' of Dumont, (miocene according to Sir C. Lyell) at Bolderberg. The specimen consists of a 
rostrum, with a portion of the base attached ; and as these parts are in some degree characteristic, I fully 
believe this specimen to be B. bisulcatus. 



BALANUS. 27 

General Appearance. — Shell (fig. 2a) conical or tubulo-conical, often rather globose ; 
walls frequently thin, either very smooth, or deeply plicated longitudinally : occasionally 
the same specimen is smooth in the upper part (fig. 2d), and strongly plicated in the 
lower. The radii in the large specimens are wide, and with their upper margins only 
slightly oblique ; in the smaller they are narrower, and much more oblique ; but in each 
case their upper margins are smooth and slightly bowed. Colour apparently originally 
nearly white, but with the alas generally, in the smaller specimens, clouded with a dark 
tint : the radii are usually striped feebly in longitudinal lines. Basal diameter of largest 
specimen *8 of an inch ; but this seems to have been an unusual size. 

Scuta : (fig. 2e) narrow, with the basal margin forming an unusually small angle with 
the occludent margin ; surface slightly convex, with lines of growth approximate, mode- 
rately prominent ; on the tergal half of the valve, two distinct rather broad furrows, with 
sometimes a third, and even a fourth, nearer to the occludent margin, extend from the 
apex down the valve, and give it a very peculiar appearance : the furrows near the tergal 
margin are the deepest. Internally (fig. 2y), the upper part of the valve is roughened 
with small points : the articular furrow is unusually wide : the articular ridge is very pro- 
minent and but little reflexed, with the lower end almost abruptly cut off: the adductor 
ridge is prominent, but short : there are small deepish pits for the rostral and lateral 
depressores. 

Terga (fig. 2/), broad, flat, with a slight narrow prominent rim along the scutal 
margin, which margin is slightly bowed. The basal margin on the carinal side of the spur 
slopes so gradually towards the spur, that the latter is barely distinct, and is very short, not 
depending nearly half its own width beneath the basi-scutal angle : the spur, also, is broad, 
namely, measured across the upper part, as broad as half the valve ; its basal end is obliquely 
rounded off on the carinal side ; it is placed close to basi-scutal angle. The carinal margin 
of the valve is just perceptibly bowed, and is formed by rectangularly upturned lines of 
growth. Internally (fig. 2/t), the upper part of the valve is rough ; the articular ridge is 
prominent ; the crests for the tergal depressores muscles are moderately well-developed. 

Parietes, not porose ; internally, the ribs are smooth, with their basal edges very finely 
or barely denticulated. The radii (as already stated) are of variable breadth ; they have 
their upper margins either very slightly or highly oblique, but always smooth and rounded: 
their sutural edges are quite smooth, or sometimes, with a strong lens, traces of transverse 
strise, representing septa, can just be discovered. The alee have their upper margins very 
oblique ; their sutural edges are, in the large specimens, quite smooth ; in the younger 
ones, plainly crenated ; the recipient furrow being clearly marked by the teeth. Basis 
plainly porose. 

Varieties. — It is certain (fig. 2d) that there are longitudinally plicated specimens of 
this species, and that the obliquity of the upper margins of the radii also varies a little ; 
nevertheless some of the deeply plicated specimens (fig. 2c) undoubtedly have a very 



28 FOSSIL CIRRIPEDIA. 

different aspect from the ordinary varieties, and do really differ in the sutural edges of the 
alee being crenated, and in the greater narrowness and obliquity of the radii ; but these 
points are all commonly variable. I have not seen any large specimens of the variety 
(fig. %c), plicatus, so as to compare them with the large specimens of the normal form, yet 
I can hardly entertain any doubt, considering their agreement in so many important 
points, that I have rightly treated these forms as mere varieties ; it is unfortunate that 
none of the specimens of the var. plicatus seen by me have had opercular valves, as their 
presence would have removed all shadow of doubt. I have given a drawing, enlarged seven 
times, of some very young shells (fig. 2d), adhering in numbers on Pecten Gerardii, which I 
believe belong to the plicated variety of our present species, but which are much too young 
to be identified with certainty. 

Affinities: this is a strongly characterised species, and nearly allied only to the 
following species, B. dolosus. The furrows on the scuta in some degree resemble those on 
the recent B. Icevis, but there is no alliance with that species. It is certain that amongst 
recent species, the chief affinity is with B. Hameri and amaryllis. 



9. Balanus dolosus. Tab. II, fig. 3a — 3d. 

B. nee parietibus, nee radiis poris perforatis ; basi poris magnis perforata, radiorum 
marginibus superioribus obliquis, lavibus ; aciebus suturalibus item leevibus ; tergi calcare 
non admodum brevi, | valvce latitudine. 

Neither walls nor radii permeated by pores ; basis permeated by large pores ; radii with 
their upper margins oblique and smooth ; sutural edges smooth ; tergum with the spur not 
very short, broad as one third of valve. 

Fossil in Red (Sutton) and Mammaliferous Crag; Mus. S. Wood, Bowerbank, Lyell, J. de C. Sowerby, 
Henslow, &c. Mammaliferous Crag, Postwick, near Norwich, Mus. Lyell. 

This species so closely resembles B. bisulcatus, both externally and in all the essential 
characters of the parietes, radii, and basis, that it is quite superfluous to describe 
again these parts. The specific characters are derived from the opercular valves, which 
present well defined distinctions, found by me constant in several specimens of both 
species. B. dolosus, like B. bisulcatus, has quite smooth and deeply plicated varieties, 
often adhering to the same univalve. The ribs on the inner surfaces of the parietes are 
remarkably prominent, as shown in the drawing (fig. 3a) of the inside of the rostrum. I 
think the upper margins of the radii are in this species rather more oblique than in 
B. bisulcatus. The sutural edges of the radii are marked by the finest strise, representing 
septa. The sutural edges of the alee are generally distinctly crenated. The basis is often 
slightly cup-formed, and very plainly porose (fig. 3b) •. its upper surface is marked by 



BAIANUS. 29 

radiating ridges : the septa between the radiating pores are themselves often in part porose, 
as was plainly the case in the specimen (fig. 3b) engraved. The orifice of the shell is 
large and elongated in its rostro-carinal axis, especially in yonng specimens. The basal 
diameter of the largest specimen is '4 of an inch. 

The scuta (fig. 3c) have no trace of the two or three longitudinal furrows so con- 
spicuous on these valves in B. bisulcatus, and which, in that species, run down to the basal 
margin from the apex of the valve, this fact showing that the furrows occur in quite young 
individuals. The whole valve is not quite so narrow as in B. bisulcatus, but otherwise 
agrees with it in shape : internally, there is hardly any difference ; but the articular furrow is 
not quite so wide : the articular ridge is very prominent, and abruptly truncated at its lower 
end : the adductor ridge is also prominent ; it here runs a little higher up the valve than 
in B. bisulcatus. The tergum (fig. 3d) differs more in the two species : the spur is not so 
broad ; measured in its upper part, it is only about one third of the entire width of the 
valve, instead of being half as wide as the valve : it is considerably longer, depending 
beneath the basi-scutal angle more than half its own width : the basal margin of the valve 
on the carinal side, does not slope so gradually into the spur; the occludent and carinal 
margins are slightly arched, as in B. bisulcatus. Internally, the surface is rough, the 
articular ridge is prominent, and the crests for the tergal depressores are well developed, — 
all as in B. bisulcatus. It is remarkable how generally the opercular valves have been 
preserved in this species in its fossil condition, as compared with most other species of the 
genus. 

It is not easy to distinguish, by external characters, the rugged varieties of this species 
from B. crenatus; indeed, the only difference is that the furrows receiving the edges of the 
radii, generally, exhibit in B. crenatus slight impressions of the septa, which are entirely 
absent in B. dolosus. By internal characters, such as the non-porose parietes, and porose 
basis, our present species widely differs from B. crenatus. 

10. Balanus tjnguiformis. Tab. II, fig. 4a — 4/. 

Balanus unguiformis. J. de C. Sowerbij (!) Mineral Conchology (sine descriptione), 

Tab. 648, fig. 1, (Jan. 1846.) 

— erisma. J. de C. Sowerby (!) lb., fig. 2. 

— perplexus. Nyst, apud D'Omalius (sine descript. vel Tab.), G6ologie de la 

Belgique, 1853. 1 

B. parietibus tenuibus, interdum ports perforatis : radiis sine poris, marginibus su- 
perioribus obliquis ; aciebus suturalibus tenuissime crenatis .• basi sine poris : tergi calcare 
angusto, obtuso. 

1 I am mucb indebted to M. Bosquet for specimens bearing this title, from Klein Spauwen, which 
certainly appear to me, as far as can be judged by the separated compartments, without the opercular valves, 
to belong to our present species. 



30 FOSSIL CIRRIPEDIA. 

Parietes thin, sometimes permeated by pores ; radii without pores, with their upper 
margins oblique ; sutural edges very finely crenated ; basis without pores. Tergum with 
the spur narrow, bluntly pointed. 

Var. erisma (fig. 43), with the walls longitudinally folded or ribbed. 

Fossil in the Eocene formations, Isle of Wight, Colwell Bay ; Hordwell ; Barton, (Chama Bed) ; Headon ; 
Bembridge ; Bergh, near Klein Spauwen, Belgium (?). Attached to various shells and wood. Mus J. de 
C. Sowerby, E. Forbes, F. Edwards, Charlsworth, T. Wright, Bowerbank, Tennant, Bosquet. 

This species, the most ancient one as yet well known in the genus, presents to the 
systematist a most unfortunate peculiarity, in the parietes being almost as often as not 
permeated by small pores : I have seen no other instance, except to a limited degree in the 
recent B.glandula, of this character being variable, and hence it must be still considered 
of high classificatory value, in so varying a genus as Balanus. Owing to the kindness of 
Mr. F. Edwards, I have seen the original specimens, excellently figured by Mr. J. de C. 
Sowerby, in the ' Mineral Conchology,' under the names of B. unguiformis and erisma, 
between which I can perceive no difference, excepting that the walls in the latter are 
longitudinally folded, — a character we know to be variable in many species. In both 
varieties, the parietes are sometimes porose and sometimes solid. The smaller specimens, 
however, figured in the ' Mineral Conchology' to the right hand of the Plate, may possibly 
be a distinct species, as I infer from the narrowness of their radii. This species is inti- 
mately allied to B. varians, a fossil from the ancient tertiary plains of Patagonia. It is also 
allied to the recent B. crenatus and glandula. 

General appearance. — Shell (fig. 4a), tubulo-conical, sometimes even sub-cylindrical: 
surface either very smooth, or slightly folded, or deeply folded so as be strongly ribbed 
longitudinally: orifice rather large, rhomboidal, narrow at the carinal end, toothed, 
but not deeply : walls rather thin and fragile : radii of moderate width, with their summits 
oblique, not quite smooth. Basal diameter of largest specimen about three quarters of an 
inch. 

Scuta (fig. 4c), with the external surface smooth : there is a trace of a furrow running 
down the valve from the apex, near to the occludent margin, and this is only worth 
mentioning from the analogous furrows in B. bisculatus. Internally (fig. 4e), the upper 
surface of the valve is roughened : the articular ridge is very prominent, and slightly 
reflexed : there is no distinct adductor ridge ; there is a slight but variable depression for 
the lateral depressor. Tergum (fig. 4d), with the longitudinal furrow shallow; spur 
moderately long, about one fourth or one fifth of the width of the valve ; placed at about 
its own width from the basi-scutal angle ; basal end bluntly pointed ; the basal margin on 
the opposite sides of the spur forms a nearly straight line ; the carinal margin has an 
extremely narrow border formed by upturned lines of growth. Internally (fig. 4/), the 
surface is roughened with little points : the articular ridge is prominent : the crests for the 
tergal depressores moderately prominent. 



BALANUS. 31 

Parietes : the longitudinal ribs on the internal surface are either feebly, or, in the 
lower part, strongly developed ; their basal ends are only just perceptibly denticulated. As 
already stated, in about half the specimens, there were no traces of parietal pores ; in the 
other half there were either distinct or obscure pores ; the pores are circular, generally of 
unequal sizes, and never large ; in the same individual they would sometimes be almost 
wholly absent in some of the compartments, and quite plain in the other compartments. 
The radii are either moderately wide or rather narrow, and have their upper margins very 
oblique, and not distinctly arched, and not quite smooth : their sutural edges are very 
finely crenated, the teeth or septa not being denticulated. The upper margins of the aloe 
are rather less oblique than those of the radii : their sutural edges are barely crenated. 
The basis is thin, and without any trace of pores ; the upper surface is sometimes furrowed 
in radiating lines. 



11. Balanus inclusus. Tab. II, fig. ba — bg. 

JB. nee parielibus, nee radiis ports perf oralis ; basi p oris perforata : testa rufo-fuscd .- 
radiis latis, marginibus superioribus aut non obliquis aut modice ; aciebus suturalibus cum 
septis plane denticulatis : scuto sine adductoris crista: tergi calcare subangusto. 

Neither walls nor radii permeated by pores ; basis porose ; shell reddish- brown ; radii 
broad, with their upper margins not oblique, or only moderately oblique ; sutural edges 
with plainly denticulated septa : scutum without an adductor ridge ; tergum with the spur 
rather narrow. 

Far. (a) (fig. 5c, 5d), with the shell elongated in its rostro-carinal axis ; basis narrow, 
clomping the stem of a zoophyte ; lateral compartments much broader than the almost linear 
rostrum, carina, and carino-lateral compartments. 

Var. {b), with rough longitudinally folded walls, and with the summits of the radii forming 
an a?igle of about 45° with the basis. 

Fossil in Coralline Crag; Sutton and Gedgrave ; attached to foliaceous Bryozoa ; Mus. S. Wood, 
Bowerbank. Far. a, Coralline Crag, Sutton, attached to cylindrical branches of corals ; Mus. S. Wood, 
Bowerbank. Far. b, attached to shells, Osnabruck, Hanover, Mus. Lyell ; Bunde, Westphalia, Mus. 
Krantz. 

My materials consist of a beautiful series of specimens in Messrs. Wood and Bower- 
bank's collections ; but unfortunately only a single young specimen had its opercular valves 
preserved. Not one specimen of the very curious variety (a) had opercular valves, yet I 
cannot feel any doubt about its being only a variety, caused by its attachment to a thin 
cylindrical branch of a coral, instead of to a foliaceous Bryozoon ; it will, however, be con- 
venient to give a separate description of this very remarkable form. With respect to 
var. (b), both sets of specimens came to me from the Continent, with the name of 



32 FOSSIL CIRRIPEDIA. 

B. stettaris, of Bronn ; but as Bronn distinctly states, that in his species the parietes are 
porose, and as such is not here the case, this cannot possibly be that species : these 
specimens did not possess their opercular valves, and therefore cannot be identified with 
certainty. 

General Appearance. — Shell conical (fig. ha, hb), with the orifice rather large, and 
rhomboidal. The surface is very smooth, except in var. (b) from the Continent, in which it 
is rugged and longitudinally folded. The colour is ochreous-brown (chiefly no doubt 
derived from the imbedding substance), tinged with red. The radii often have a much 
darker and more distinct red tint ; they are sometimes longitudinally striped with dirty 
white. The radii are broad, with their summits straight, and very slightly oblique ; in 
var. b, however, they slope at an angle of about 45°. Basal diameter of largest specimens 
•6 of an inch ; but this is an unusual size. 

Scuta, with the growth ridges little prominent. Internally (fig. 5/, from a young 
individual) the articular ridge is moderately prominent, with its lower end very obliquely 
rounded off; there is no adductor ridge ; there is a minute pit for the lateral depressor 
muscle. Terga, with a slight longitudinal depression extending down to the spur ; spur 
short, with its lower end almost square or truncated, about one fourth of width of valve, 
and placed at about half its own width from the basi-scutal angle. Internally (fig. hg) 
the articular ridge is prominent ; the crests for the tergal depressores are feebly developed. 

Parietes, moderately thick and generally strongly ribbed internally, without parietal 
pores. Radii, wide, with their upper margins straight, not smooth or rounded, and very 
slightly (or, in var. b, moderately) oblique ; their sutural edges have well-developed septa, 
which are denticulated : the interspaces between the septa are filled up solidly. The alee 
have their upper margins oblique: they are only slightly, and sometimes not at all, added 
to above the level of the opercular membrane : their sutural edges are smooth. The basis 
is thin, but plainly porose. 

Var. {a) (fig. 5c, hd). — With respect to this remarkable variety, any one would at first 
think it specifically distinct. The shell is much compressed, or elongated in the rostro- 
carinal axis, sometimes to a great degree; I have seen a specimen - 25 of an inch in this 
axis, and only *1 in its broadest part ; but this is a very unusual degree of elongation. The 
most remarkable character is the extraordinary narrowness of the carina, of the carino-lateral 
compartments, and of the rostrum, compared with the great breadth, especially along the 
basal margin (fig. hd), of the lateral compartments. The radii are of unusual breadth. The 
tips of the rostrum and of the lateral compartments are a little arched in, tending to make 
the shell somewhat globular. The true basis is extremely narrow (fig. hd) : it is deeply 
grooved, from clasping the thin, cylindrical stem of the coral to which it has adhered ; and 
I have seen specimens in which the opposite edges of the groove had met, a tube having 
been thus actually formed. From the grooved basis, and from the elongation of the shell 
in the rostro-carinal axis, this variety presents so close a general resemblance to Balanus 
calceolus, and its allies, that I have seen it in a collection arranged on the same tablet with 



ACASTA. 33 

a fossil specimen of B. calceolus. Notwithstanding the above several strongly-marked 
characters, by which this variety differs from the ordinary form, there is a resemblance in 
colour and aspect, which, though difficult to be described, made me from the first suspect 
that the two were specifically identical. In no point of real structure is there any difference, 
excepting that, perhaps, the pores in the basis are here rather smaller ; but this might arise 
from the little development of the peculiar basis. Having come to this conclusion, I was 
interested by finding a specimen (fig. be) in Mr. Wood's collection, which had originally 
fixed itself (judging from the form of the basis) on a thick cylindrical stem, but which had sub- 
sequently grown on to an adjoining flat surface ; consequently, one side of the shell presented 
all the peculiar characters of the present variety, but not strongly pronounced, whereas the 
other side, at the rostral end, was undistinguishable from the ordinary form. The unequal 
development of the rostrum on the two sides was very striking, and clearly showed how 
great an effect could be produced by the nature of the surface of attachment. 

This singular variety cannot be considered accidental, in the sense in which this term 
may be applied in some cases : the pupa evidently fixes itself intentionally, in a certain 
definite position, on the branch of the coral (when a branch is chosen), exactly as in the case 
of Balanus calceolus, or Scapettum vulgar -e, — species which always live attached to branches. 
But when other Balani occasionally fix themselves on branched corals, their position seems 
to be accidental and unsymmetrical; thus among the symmetrically elongated specimens of the 
present species, I found one specimen of Balanus bisulcatus, which had evidently been 
attached in an almost transverse position to a branch, and had thus become much distorted ; 
so, again, I have seen specimens of the recent B. amargltis attached irregularly to a Gorgonia, 
in the midst of the symmetrically elongated shells of Balanus navicula, an ally of B. calceolus. 

This variety does not seem to attain so large a size as the ordinary form. 

Affinities. — This species is allied to B. unguiformis and B. varians, but is perhaps more 
nearly related to the recent B. allium, an inhabitant of the Barrier Reef of Australia. The 
longitudinally folded variety (b) can hardly be distinguished by external aspect, or even by 
the opercular valves, from B. crenatus ; but when the shell is disarticulated, the porose 
walls and non-porose basis of B. crenatus, allow of no mistake in the diagnosis of the 
two species. 



Sub-Genus — Acasta. 
Acasta. Leach. Journal de Physique, torn. Ixxxv, 1817. 

Valvee testa 6; 2 j arietes et basis non porosa ; basis calcarea, cyathiformis, non elongaia. 
Vulvae operculares inter se articulates, sub triangulares. Spongiis, ant rarb Isidis cortici, 
affix a. 

Compartments six ; parietes and basis non-porose : basis ealcareous, cup-formed, not 

5 



34 FOSSIL CIRRIPEDIA. 

elongated, attached to Sponges, or rarely to the bark of Isis : scutum and tergurn articulated 
together, subtriangular. 

Under the last genus, I have made a few remarks on the close affinities of this sub-genus 
to Balanus, and have given my reasons for retaining it, so that I need not here 
repeat them. 



Acasta unuulata. Tab. II, fig. 6a — 6f. 

A. testa, ad speciem, ut in " A. spongites," sed majore : scuto externe striis longitudi- 
nalibus, sape binis, signato, sulcis intermediis latioribus .- tergi calcare, pane \ valvce 
latitudine. 

Shell, apparently, as in A. spongites, but larger : scutum marked by longitudinal ridges, 
often in pairs, with the intermediate furrows rather wide : spur of tergum nearly half as 
wide as valve. 

Fossil in Coralline Crag (Sutton), Mus. S. Wood, Bowerbank. 

I owe to Mr. Wood the inspection of a fine suite of valves, which, though separate, I 
have no reason to doubt have all been rightly attributed to the same species. Owing to the 
shell never having been found entire, its general shape is not known, and, what is of more 
consequence, the relative proportional width of the parietes of the carino-lateral compart- 
ment is unknown. I have (but with doubt) given it a distinct specific name, owing to the 
peculiar character of the furrows on the scuta, and to the large size of the whole shell. In 
its other characters it comes nearest to A. spongites, excepting in the spur of the tergum, 
which resembles that of A. sulcata. 

The external surfaces of the compartments appear generally to have been smooth ; but 
in several specimens they are studded with the sharp shelly points so common in the 
genus. A rostrum (Tab. II, fig. 6a), and lateral compartment (fig. 6b), have been figured. 
The radii are not wide. The basis (fig. 6c) is cup-formed : its edge is either quite smooth, 
or is very finely crenated. The basis is sometimes quite irregularly perforated, as in the case 
of several recent species, by numerous minute orifices, which, when the animal was 
alive, no doubt were covered by membrane. Internally the parietes are feebly ribbed, as 
in A. spongites. Judging from the dimensions of the separated valves, this species must 
have equalled and perhaps exceeded in size the largest living species, namely, A.glans, from 
Australia. Hence w r e may infer, that the basal diameter probably exceeded '55 of an 
inch : I may add, that the largest European specimens of A. spongites, from Naples and 
Portugal, are only '3 of an inch in basal diameter. 

Scuta (fig. 6e). — These seem to resemble the scuta of A. spongites in all respects, except 



PYRGOMA. 35 

in the external longitudinal ridges standing much further apart, and, consequently, in the 
farrows being much wider : each ridge is generally double. Although there is a good deal 
of variability in the character of these ridges in A. undulata, and likewise in A. spongites, I 
have not seen any form intermediate between them. It must, however, be confessed, that 
this is an extremely variable character in many sessile cirripedes. Internally the scutum 
(fig. 6d) is chiefly characterised by the absence of characters, that is, by the slightness of 
the pits for the muscles, and the little prominence of the articular ridge. In the tergum 
(fig. 6/), the spur is about half the width of the whole valve, and therefore rather wider 
than in A. spongites. 



Genus — Pyrgoma. 

Pyrgoma. Leach. Journal de Physique, torn. 85, 1817. 
Boscia. Ferussac. Diet. Classique d'Hist. Naturelle, 1822. 
Sayignium. Leach. Zoological Journal, vol. ii, July, 1825. 
Megatrema. lb. lb. 

Adna. lb. lb. 

Daracia. J. E. Gray. Annals of Phil, (new series), August, 1825. 
Creusia. Be Blainville. Diet. Sc. Nat., PI. 116, 1816-30. 
Nobia. G. B. Sowerby , junr . Conchological Manual, 1 1839. 

Valves testes in unam confluente : basis cyatldformis aut subcylindrica, coraliis affixa : 
vafoce opercidares inter se articidatce. 

Shell formed of a single piece : basis cup-formed, or subcylindrical, attached to corals : 
scutum and tergum articulated together. 

This genus can at once be recognised by the shell consisting of a single piece without 
sutures, whether viewed externally or internally, and by the cup-shaped basis, attached and 
often imbedded in corals. The one species, P. Anglicum, found both recent and fossil, 
together with a closely allied recent species, P. Stokesii, in all the characters derived from 
the opercular valves, closely resemble Balanus and other ordinary forms, and for this very 
reason they have some slight claims to be generically separated from the other species 'of 
Pyrgoma ; for in these latter, the opercular valves seem to have broken loose from all law, 
presenting a greater diversity in character than do all the other species of Balaninae and 
and Chthainalinse taken together. 

1 The name, Nobia, is given in this work on the authority of Leach, but this must be a mistake, pro- 
bably caused by some MS. name, (that fertile source of error in nomenclature), having been used. 



36 FOSSIL CIRRIPEDIA. 



Pyrgoma Anglicum. Tab. II, fig. la — 7c. 

Pyrgoma Anglica. G. B. Sowerby. Genera of Recent and Fossil Shells, fig. 7, 

No. 18, Sept. 1823 (sine descript.). 
Megatrema (Adna) Anglica. J. E. Gray. Annals of Philosoph. (new series), vol. x, 

Aug. 1825. 
Pyrgoma sulcatum. Philippi. Enumeratio Molluscorurn Sicilise, Tab. 12, fig. 24, 
(1836). 
— angltca. Brown. Illustrations of Conchology, (2d edit., 1844), Tab. 53, 

fig. 27—29. 

P. testa abrupte conicd, purjmreo -rubra ,• orificio ovato, angusto ; basi porosd, plerumque 
e coralio exserta : scuto et tergo subtriangularibus. 

Shell steeply conical, purplish red : orifice oval, narrow : basis permeated by pores, 
generally exserted out of the coral : scutum and tergum sub-triangular. 

Fossil in the Coralline Crag (Ramsholt) Mus. S. Wood. 

Recent on the south coast of England and of Ireland, (12 to 45 fathoms, Forbes and MacAndrew) ; 
Sicily ; Madeira ; St. Jago, Cape de Verde Islands ; generally attached to the edge of the cup of a 
Caryophyllia, in deep water, but at St. Jago within the tidal limits. 

I have considered this fossil as identical with the recent species, but, as may be seen 
from the following description, it presents several slight differences; yet they are snch that 
I dare not found a new species on only a few specimens thus characterised. 

The shell is steeply conical, slightly compressed, with the lower part having rounded, 
approximate, radiating ribs ; these ribs seem to be more prominent in the fossil than in 
the recent specimens. Colour dull purplish-red. Orifice oval, small, and narrow. The 
basis is not deeply conical, and occasionally is even flat: in the Crag specimens it is almost 
wholly imbedded in the coral to which it is attached ; but in recent specimens it is gene- 
rally exserted. Externally the basis is furnished with ribs corresponding with those on the 
shell. The largest recent specimens which I have seen, from St. Jago, was '22 of an inch 
in basal diameter ; but some few of the British specimens are nearly as large, and one of 
the fossils from the Coralline Crag a very little larger. 

The scuta and terga are of the ordinary shape of these valves in Balanus and its 
allies. Scutum (fig. 7b) triangular, with the basal margin a little curved and protuberant ; 
adductor and articular ridges distinct from each other, moderately prominent; there is 
a small hollow for the lateral depressor muscle : in the fossils, the adductor ridge (as 
figured) is more distinct from the articular ridge, and consequently the cavity for the 
lateral depressor muscle is wider and less deep that in recent specimens. Tergum, I have not 
seen a fossil specimen, but have figured a recent valve (7c) ; it is triangular, with the spur rather 
narrow, moderately long, placed near, but not confluent with, the basi-scutal angle of the 



CORONULA. 37 

valve. The basal margin forms an angle rather above a right angle with the spur. Inter- 
nally the articular ridge and crests for the depressor muscles, feebly developed. 

Internal Structure of the Shell and Basis. — Internally, the shell is ribbed, more or less 
prominently. The lower edge of the sheath, which is reddish, and extends far down the 
walls, seems always to project freely. In several specimens there were on each side, at 
the carinal end of the shell, a trace apparently of a suture, which could be perceived only 
on the sheath. The basis appears always to be permeated by minute tubes or pores, 
though these are sometimes rather difficult to be seen. 

Michelotti, in the 'Bulletin Soc. Geolog.' Tom. 10, p. 141, has named, but not 
described, a species, viz., Fyrgoma utidata, from the northern Italian Tertiary Strata. 

Genus — Coronula. 

Coronula. Lamarck. Annales du Museum, torn, i, (1802). 

Diadema. Schumacher. Essai d'un Nouveau Syst., &c, 1817. 

Cetopirus (sed non Coronula). Ranzani. Mernoire di Storia Naturale, (1820). 

Polylepas. J. E. Gray, (Klein). Annals of Philosophy, (new series), vol. 10, 1825. 

Valva tastes 6, cequali latitudine ; parietes tenues, profunde plicati, plicis cavitates 
infra solum apertas efficientibus ; valva opercidares non inter se articulates, orificio testae 
multo minores : basis membranacea. Cetaceis afjixa. 

Compartments six, of equal sizes : walls thin, deeply folded, with the folds forming 
cavities, open only on the under side of the shell : opercular valves much smaller than the 
orifice of the shell ; when both present not articulated together : basis membranous. 
Attached to Cetaceans. 

The structure of the shell of Coronula is complicated, and has been generally quite mis- 
understood. Without a long description and several figures it would be impossible to 
give a true idea of its singular structure ; but, in order to make the following description 
at all intelligible, I must make a few remarks. The wall of each compartment, and there- 
fore of the whole shell, is extremely thin ; but strength is gained by its being folded in a 
very complicated manner, as may be seen in the rostral compartment, Tab. II, fig. 8b, 
by tracing the wall e to e , to e"; the folds at their outer ends are elongated into transverse 
loops, the extremities of which touch each other ; consequently, what appears to be the out- 
side of the shell consists only of a portion of the wall, namely, the outsides of the transverse 
circumferential loops, together with the radii. These loops appear externally like much 
flattened longitudinal broad ribs. On the other hand, the inside of the shell, in which the 
body is lodged, consists of the inner ends of the folded walls, lined by the sheath, and by 
the alse. The basal edges of the folded walls, in the line of the ray of the circular shell, 
are oblique ; the outer ends, or transverse circumferential loops, having grown downwards 
at a greater rate than the inner ends. Between each fold of the walls, there is a flattened 



38 FOSSIL CIRRTPEDIA. 

cavity, open at the bottom of the shell, and running up to the apex ■ these cavities are quite 
external to the cirripede, and are occupied by the epidermis of the whale to which the 
Coronula is attached : homologically they are only deep longitudinal furrows, and they would 
still have been furrows, had not the transversely elongated ends of the folds, i. e. } the 
circumferential loops, in all cases, after early growth, grown into close contact. The ends of 
these loops are generally locked together by rows of minute teeth. In all the species, 
when young, the wall of each compartment is folded three times, and therefore the whole 
shell has eighteen folds. 

The radii, normally, are only part of the wall, modified by growing against an opposed 
compartment ; and hence the radius in Coronula would have been extremely thin, like the 
wall, and the sutures between the six compartments excessively weak, had not the radii 
been specially thickened by numerous sinuous denticulated plates, springing from the inner 
lamina of the true radius, and running downwards, attached to the folded wall of the com- 
partment to which the radius belongs, and with their free edges pressed against the folded 
wall of the opposed compartment. Hence the radii may be said to be compound. For 
the sake of strengthening the sutures, the alas, also, are very unusually thick : but, notwith- 
standing their thickness and the thickness of the compound radii, owing to the depth of 
the folds of wall, they are separated from each other by a considerable space, and the alas, 
instead of resting in chief part, as they should do, on the inner lamina of the radius, have 
to rest on special plates, developed apparently from the sheath. In the upper part of the 
shell, between the special plates on which the alas rest, and the compound radii, there are 
in two of the three recent species, open chambers, six in number, occupied by the ovarian 
caeca ; but in the fossil C. barbara these chambers are almost filled up solidly by shell. I 
hope that the terms used in the following description may be now in some partial degree 
rendered intelligible. 



Coronula Barbara. Tab. II, fig. 80 — Be. 

CoRONULiTES diadema (?) Parkinson. Organic Remains (1811), vol. iii, p. 240, 

pi. 16, fig. 19. 

C. testa {probabilitef) coroniformi, costis longitudinalibus convexis, aciebus earum cre- 
natis, superflcie interna et externa cristis transversis asperd ; radiis modice crassis ; spatio 
inter radios et alas solide impleto. 

Shell (probably) crown-shaped, with longitudinal convex ribs, having their edges 
crenated, and their surfaces rugged, both externally and internally, with transverse ridges : 
radii moderately thick ; the space between the radii and the alae solidly filled up. 

Fossil in Red Crag, (Bawdsey and Sutton) ; Mus. S. Wood and Geological Society. 






CORONULA. 39 

This species, though closely allied to C. diadema and easily confounded with it, I have 
no doubt is distinct. I owe to the kindness of the Rev. Mr. Image an examination of the 
original specimen figured by Parkinson ; and in Mr. Stutchbury's collection there is a 
similar and more perfect specimen ; both of these resemble C. diadema in general form, 
but have been too much worn to be positively identified. The following description is 
drawn up from some compartments collected by Mr. Searles Wood, belonging certainly to 
three and probably to four individuals, one of which was young; as these specimens agree 
in all essential respects, I feel pretty confident that the characters, by which the present 
species differ from C. diadema, are of specific value. 

Structure of Shell. — The longitudinal ribs on each compartment (i. e. the circumferen- 
tial transverse loops), are convex and prominent, as in C. diadema, but they are crossed by 
more prominent ridges of growth (fig. 8a, 8e) than even in the roughest varieties of that 
species, so that the surface of the shell is more rugged. In the three recent species — viz., 
C. diadema, balcenaris, and regince, the surface of the wall all round the cavities oc- 
cupied by the whale's skin, is striated only by very fine longitudinal lines ; but here, the 
outer portion, or that (fig. 8c/) formed by the transverse loops, is crossed by transverse 
ridges of growth, like, but less prominent, than those on the external surface of the shell. 
The minute teeth, along the lines of junction between the transverse loops, are here less 
regular, and can hardly be said to exist ; for the two edges are locked together by what may 
be more strictly called minute zig-zag ridges (fig. 8d, 8<?), than teeth. The exact number of the 
circumferential plications (fig. 8b) in the wall of the shell is variable, in the same manner as 
in the three recent species. In the rostrum which has been figured (86, enlarged twice 
its natural size), there is a peculiarity, probably accidental, which I have seen in no 
other specimen — namely, that one of the transverse circumferential loops at the end of one 
of the original folds of the wall, has ceased to be added to, and therefore may be seen 
(rather on the right hand of the middle of the figure) to terminate in one of the cavities 
between two adjoining folds. The sutural edges of the compound radii (d, fig. 8b) 
are about as thick as, or rather thicker than, in C. diadema; for in the middle part 
they do not reach to the sheath by about half the thickness of the compartment. In the 
same manner as in C. diadema and regince, each ala here rests, not on the internal surface 
(as in C. balcenaris, and in all other Balanidse) of the radius, but on a special plate (c, fig. 
8b, 8c) ; but in C. barbara, instead of there being a deep chamber, running up to the apex 
of the compartment, between the radius and the special plate, this part is filled up almost 
entirely by solid shell. Although the extent to which this chamber is filled up varies a little, 
and although its depth varies a little in C. diadema, yet there is a marked difference between 
the specimens of this latter species, in which the chamber is most filled up, and those of 
C. barbara, in which it is least filled up. The alse are thick, as in C. diadema, and their 
sutural edges have a central ridge, sending off on both sides sinuous crests. The basal 
margins of the alas are not short compared with their upper margins, and therefore the 
whole ala is not wedge-formed (fig. 8c); and in this rather important respect C. barbara 



40 FOSSIL CIRRIPEDIA. 

resembles C. balanaris, and differs from C. diadema. The lower edge of the sheath does 
not seem to have projected freely, — in this respect, also, resembling C. balanaris. From 
the basal margin of the alse not being narrow, and from the inner ends of the folded walls 
being, as it would appear, also broad, I have little doubt that the cavity in which the 
animal's body was lodged, resembled in shape that in C. balanaris, the membranous basis 
being larger than the orifice of the shell. 

Opercular valves unknown. 

Summary. — This species is more nearly related to C. diadema than to the others ; but 
in some points, just specified, it resembles C. balanaris. The characters by which it 
differs from all the species are, firstly, the more prominent transverse ridges on the 
external surface of the shell, and more especially on the surfaces bounding the outer sides 
of the cavities occupied by the epidermis of the whale. Secondly, the character of the 
teeth, or rather ridges, along the lines of junction between the transverse loops. And, 
thirdly, the spaces between the radii and the special plates on which the alas rest, being 
solidly filled up. 

The Coronula bifida is an Italian tertiary species, so named by Bronn, in his " Italiens 
Tertiar-Gebilde" (1S31), p. 126. It is very possible that this may be identical with 
C. barbara, but Bronn does not seem to have been aware of the absolute necessity of 
giving minute details in his descriptions of fossil cirripedes. The chief character of 
C. bifida is thus given : — " Eine tiefe Furcle oder Spalte theilt die Laugenrippe von oben 
herab bis zur Halfte, welche bei der sonst ahnlichen C. diadema entweder ganz fehlt, oder nur 
zuweilen kurz augedeutet ist." Had it been stated that the longitudinal ribs were divided 
from the middle down to the base, instead of from the top to the middle, the description 
would have been intelligible to me, though the character thus afforded would not have 
been of specific value, as this dividing of the ribs occasionally occurs in all four species, 
and is produced by the formation of new folds in the walls. 



Family — Verrucid^e. 

Cirripedia sine pedunculo : scuta et terga, musculis depressoribus non instructa, ex uno 
latere tantum mobilia, ex altero cum carina et rostro in testam asymmetricam immobiliter 
conjunct a. 

Cirripedia without a peduncle : scuta and terga, not furnished with depressor muscles, 
moveable only on one side, on the other side united immoveably with the rostrum and 
carina into an unsymmetrical shell. 



VERRUCA. 41 



Genus — Verruca. 

Verruca. 1 Schumacher. Essai d'un Nouveau Syst. Class., 1817. 

Clysia. Leach. Journal de Physique, torn. 85, July, 1817; Clisia, Leach, Encyclop. 

Brit. Suppl., vol. iii, 1824; Clitia, G. B. Sowerby, Genera of Recent and 

Fossil Shells. 
Creusia. Lamarck. Animaux sans Vertebres, 1818. 
Ochthosia. Ranzani. Memoire di Storia Nat., 1820. 
Lepas et Balanus Auctorum. 

The family of Verrucidae includes only the above single genus ; but it has, I think, as 
good a claim to be considered a distinct Family as either the Balanidae or Lepadidae, that 
is, either the Sessile or Pedunculated Cirripedes. The two latter Families differ from each 
other almost exclusively in the nature of the shell or external covering, and in the 
muscles moving the different portions of it : now Verruca has a very peculiar shell, 
destitute of all muscles, excepting the adductor scutorum, and composed of only six 
valves, and these are so unequally developed, that the longitudinal dorso-ventral plane of 
the body comes to lie nearly parallel to the surface of attachment, instead of at right angles 
to it. Upon the whole, the Verrucidae are nearly equally related to the Lepadidae and 
Balanidae ; but certainly nearer to the Lepadidae, than to the sub-family Balaninae or typical 
sessile cirripedes ; though, on the other hand, if compelled to place Verruca in one of 
these two Families, I should place it amongst the Chthamalinae, the other sub-family of 
Balanidae. The distinctness of Verruca, though in appearance a sessile cirripede, from 
the Balanidae or true sessile cirripedes, is interesting, inasmuch as no member of this latter 
Family has hitherto been found fossil in any Secondary Deposit, whereas Verruca ranges 
from the present day to the upper beds of the Chalk near Norwich, and in Belgium ; 
being likewise found in the Glacial Deposits, in the Red and Coralline Crags of England, 
and in an ancient tertiary formation of Patagonia. 

The shell of Verruca has generally been quite misunderstood: it consists, as already 
stated, of six valves ; and these can be proved (as I have shown in my volume published 
by the Ray Society), by tracing the development of the young shell, to consist of a rostrum 
and carina, unequally developed on their two sides, — of a scutum and tergum in their 
normal and moveable condition, — and, lastly, of the scutum and tergum on the opposite 
side, most singularly modified, immoveably articulated to the rostrum and carina, forming 
together with them a shell, which is firmly united to the basal membrane, and so to the 
surface of attachment. It can be shown that the very remarkable modification and 

1 According to Bock, in the ' Naturforscher' of 1778, this term was used by Rumph for a Chelonobia, 
but as it was before the adoption of the binomial nomenclature, according to the Rules, it may be passed 
over, and does not interfere with the priority of Schumacher. 

6 



42 FOSSIL CIRRIPEDIA. 

enlargement of the fixed scutum and tergum, is due to the development of a single small 
portion in each valve, namely, the lower ridge of the articular ridges by which these 
valves are united together. It is very remarkable that in all the species it seems to be a 
matter of chance, whether the right or left hand valves undergo this singular modification ; 
consequently, of every valve it is equally likely to find a right-hand or left-hand specimen ; 
and these, though exactly alike, except in being reversed, or in coming from opposite sides 
of the body, yet, from this very circumstance, and from the fixed valves being of very 
irregular shapes, are rather perplexing to identify. This short description will, I hope, 
suffice to make the following descriptions intelligible. 



1. Verruca Stromia. Tab. II, fig. da, 9 b. 

Lepas Stromia. O. Mutter. Zoolog. Dan. Prod., No. 3025, 1776. 

— — lb. Zoolog. Dan., vol. iii, Tab. 94, 1789. 

— striata. Pennant. British Zoology, vol. iv, Tab. 38, fig. 7, 1777. 

Die warzenformige meereichel. Spengler. Schriften der Berlin. Gesell., 1 B., 

Tab. 5, fig. 1—3, 1780. 
Lepas verruca. Spengler. Skrifter af Naturhist. Selskabet, 1 B., 1790. 

— — et Stromia. Gmelin. Syst. Nat., 1789. 

Balanus veruca. Bruguicre. Encyclop. Meth., 1789; Clisia verrucosa, Deshayes, in 
Tab. 
— intertextus. Pulteney. Catalogue of Shells of Dorsetshire, 1799. 
Lepas striatus. Montagu. Test. Brit., 1803. 

— verruca. Wood's General Conchology, PI. 9, fig. 5, 1815. 
Verruca Stromii. Schumacher. Essai d'un Nouveau Syst. Class., 1817. 
Creusia Stromia et verruca. Lamarck. Animaux sans Vertebres, 1818. 
Ochthosia Stroemia. Ranzani. Memoire di Storia Nat., 1820. 
Clisia striata. Leach. Encyclop. Brit. Suppl., vol. iii (sine descript.), 1824. 
Clitia verruca. G. B. Sowerby. Genera of Recent and Fossil Shells, Plate. 
Verruca Stromii. J.E. Gray. Annals of Philosophy (new series), vol. x, Aug. 1825. 

V. scuto n/obili, crista articulari inferiore dimidiam brevis crista articularis superioris 
latitudinem non aquante : testa plerumque lonyitudinaliter sulcata. 

Moveable scutum, with the lower articular ridge not half as broad as the short upper 
articular ridge : shell generally ribbed longitudinally. 

Fossil in Glacial deposits of Scotland, Mus. Lyell ; Red Crag (Walton, Essex), Coralline Crag (Sutton), 
Mus. S. V. Wood. 

Recent on the shores of Great Britain and Ireland ; Shetland Islands ; Denmark ; Iceland ; shores 
of northern Europe ; Red Sea. Attached to shells, laminarise, rocks, crabs, and floating bark, from low 
tidal mark to fifty or ninety fathoms. 

I have seen a perfect specimen of this species from the Glacial deposits of Scotland, and 



VERRUGA. 43 

separated valves from the Red and Coralline Crags, collected by Mr. S. Wood. The 
moveable opercular valves have not been discovered ; and these are certainly much the 
most important parts for the diagnosis of the species ; but the other valves are tolerably 
perfect, and are undistinguishable from recent specimens of V. Stromia ; therefore, I have 
ventured, with some hesitation, thus to name these specimens. The fossil specimens all 
belong to the common variety, having its shell longitudinally ribbed, a character not 
observed in. the four other species of the genus. As an aid to collectors in the Crag, I have 
thought it would be more serviceable to give a drawing (fig. 9 a), from a recent specimen, 
of all the six valves, separated, but in as nearly as possible their proper relative positions, 
and likewise of the under side of the fixed scutum and tergum, than to give fac-similes of 
such valves, in themselves not perfectly characteristic, which have as yet been discovered 
fossil. 

It should be borne in mind, that of the six valves of which figures are here given, it is 
just as likely that reversed specimens from the opposite side of the body should be found, 
as these which represent valves taken from a specimen in which the left-hand scutum and 
tergum were fixed and formed part of the shell. 



2. Verruca prisca. Tab. II, fig. 10a — 10c. 

Verruca prisca. Bosquet. Monographie des Crustaces fossiles du Terrain Cret. de 
Limbourg, Tab. 1, fig. 1—6; 1853. 

V. testa Icevi ; scuti mobilis crista articulari inferiore aliquanto latiore quam 
superior. 

Shell smooth : moveable scutum, with the lower articular ridge somewhat broader than 
the upper articular ridge. 

Fossil in Chalk, Norwich, Mus. J. de C. Sowerby : ' Systeme Senonien et Maestrichtien,' Belgium, 
Mus. Bosquet. 

M. Bosquet has admirably figured and described the several separated valves belonging 
to this species, and I owe to his great kindness an examination of some of them. In Mr. 
J. de C Sowerby's collection, also, there is a single specimen (fig, 10a), attached to a 
Mollusc, with the four valves of the shell united together, but without the two moveable 
opercular valves ; it cannot be positively asserted that this is the same species with that 
of M. Bosquet, but such probably is the case. The opercular valves (fig. L03, 10c) are 
necessarily figured from Belgian specimens. It is the English specimen to which I 
alluded in the Introduction to my 'Monograph on Fossil Lepadidse.' This species of 
Verruca is interesting, from being the only known Secondary one, but in itself it is a 



44 FOSSIL CIRRIPEDTA. 

very poorly characterised form, and I can point out no important character in the shell 
by which it can be recognised. The rostrum and carina, which are of nearly equal sizes, 
are locked together, and likewise to the fixed scutum and tergum, by the usual interfold- 
ing plates ; the plates between these latter valves seem to have been less developed in 
M. Bosquet's specimen than in the English. The fixed scutum has a large adductor plate, 
which seems to have been chipped in M. Bosquet's specimen ; this valve and the fixed 
tergum in all essential respects resemble the same valves in V. Stromia. The surface of 
the shell is very smooth. 

The moveable scutum has its occludent margin considerably arched : the lower articular 
ridge is broader than the upper ridge, in which respect it resembles the same valve in V. laevi- 
gata, a South American form, but the whole valve is not so broad as in that species. There is 
no adductor ridge on the under surface. The moveable tergum has its upper articular 
ridge narrow, and slightly produced into a point on the scutal margin : in this latter 
respect this species, also, resembles V. lavigata, but differs from it in the whole valve not 
being so broad in proportion to its height. 



m 



INDEX 



TO 



MONOGRAPH ON FOSSIL BALANIDiE. 



in- 



C. DARWIN, M.A., F.E.S., Ac. 



N.B.— The 


names in Italics are Synonyms. 




■ 


PAGE 1 




PAGE 


Acasta undulata . 


34 


Balanus spongicola 


16 


Adna . . . . . 


35 


,, sulcatinus 


26 


Balanus angulosus . 


21 


,, sulcatus . 


21 


„ arctica pateUiformis 


21 


,, tesselatus . 


21 


,, bisulcatus 


26 


,, tintinnabulum 


13 


,, calceolus . 


15 


,, tulipa 


13,2.5 


,, candidus . 


25 


„ Uddevallensis 


25 


,, clavatus . 


23 


., unguiformis 


29 


,, concavus . 


17 


,, verruca 


42 


„ crassus 


13 


,, zonarius . 


17 


„ crenatus . 


23 


Boseia .... 


35 


,, cylindraceus 


17 


Cetopirus . 


37 


„ dolosus 


28 


Chirona .... 


12 


,, d'Orbignii 


13 


Clitia verruca 


42 


,, elongatus 


23 


Clysia .... 


41 


., erisma 


29 


Conopea ovata . . 


15 


,, geniculatus 


. 21 


Coronula barbara . 


38 


,, glacialis . 


. 23 


Coronulites diadem « 


38 


., Hameri 


. 24 


Creusia .... 


35, 42 


., inclusus 


. 31 


Daracia .... 


35 


intertextus 


. 42 


Diadema .... 


37 


,, miser 


. 17 


Lepas balanus 


21 


„ perplexus . 


. 29 


„ borealis 


23 


,, porcatus 


. 21 


,, calceolus 


15 


,, pustular is . 


. 17 


,, costata 


21 


„ rugosus 


23 


„ crispata 


13 


,, Scoticus . 


. 21 


„ foliacea 


23,24 



INDEX. 





PASE 








PAGE 


Lepas Hameri 


. 24 


Megatrema . . . .35 


,, porcata 


. 13 


Messula . 






12 


, , Scotica 


. 21 


Nobia 






35 


spinosa 




Ochthosia . 






41 


,, striata 


. 42 


Polylepas . 






37 


„ stromia 


. 42 


Pyrgoma Anglicum 






36 


,, tintinnabulum 


13,17 


Pyrgoma sulcatum . 






36 


,, tulipa 


24 


Savignium . 






35 


,. tulipa alba . 


. . . 24 


Verruca prisca 






43 


verruca 


• 42 1 


„ stromia 






42 



TAB. I. 

Fig. la, Balanus tintinnabulum, nat. size; small specimen. 

lb, ,, „ rostrum, internal view of, large specimen, nat. size. 

Ic, Id, ,, ,, scutum and tergum, internal views of, from recent spe- 

cimens, the opercular valves not having been found 
fossil. 
Fig. 2a, Balanus calceolus, engraved from a recent specimen, the single fossil having been 

injured during examination. 
2b, 2c, „ „ scutum and tergum, internal views. 

2c?, Spur of tergum, variety : all from recent specimens, the opercular valves not 
having been found fossil. 
Fig. 3a, 3b, Balanus spongicola, scutum and tergum, external views. 
3c, 3d, „ ,, „ „ internal views. 

3c, ,, ,, shell, enlarged from a recent specimen, the single fossil 

being young, and having been injured by examination. 
Fig. 4a, Balanus concavus, shell (Coralline Crag specimen). 

4b, „ ,, internal view of part of the carina (to the left hand), of the ca- 

rino-lateral compartment, and of part of the lateral compart- 
ment, showing the raised borders (r) on the rostral sides of 
the sutures in the sheath (Italian Tertiary specimen). 
4c, „ „ shell, var., with longitudinal ridges or ribs not prominent 

(Coralline Crag). 
4c?, „ ,, smooth var. (Piedmont specimen). 

4e, „ ,, scutum, external view (Coralline Crag). 

4/, „ ,, „ internal view do. 

4g, „ „ tergum, external view do. 

4h, ,, ,, scutum, internal view (Maryland, U.S.). 

4i, „ „ tergum, internal view do. 

4k, „ „ ,, external view do. 

4/, „ ,, scutum, external view (Portugal fossil, and recent). 

4m, „ „ tergum, external view do. do 

4n, „ „ scutum, internal view do. do. 

4o, ,, „ tergum, external view, very large specimen (Turin). 

4p, ,, „ scutum, small portion, at the rostral corner, highly magnified, 

very large specimen (Turin). 
Fig. da, Balanus porcatus, shell, nat. size (Red Crag). 



FOSSIL CIRRIPEDIA. 

bb, Balanus porcatus, small portion of basal margin of wall, much enlarged. 
5c, „ „ portion of basal plate, much enlarged. 

bd, be, bf, bg, Balanus porcatus, scutum and tergum, external and internal views ; 

engraved from recent specimens, the opercular 
valves not having been positively found fossil. 
Fig. 6a, Balanus crenatus, twice nat. size (Red Crag). 

Qb, „ ,, small portion of basal margin of wall, much enlarged. 

Qc, „ „ portion of basal plate, of unusual thickness, much enlarged. 

6d, Qe, „ „ scutum and tergum, external views from recent specimens. 

6/, Qg, „ „ ,, „ war., internal views from recent specimens. 

Fig. 7a, 76, Balanus Hameri, scutum and tergum, external views engraved from Canada 

glacial specimens, the opercular valves not having been 
found in Great Britain. 
7c, 7d, „ „ scutum and tergum, internal views of very large and old 

specimens (Canada andUddevalla). 



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TAB. II. 

Fig. la, Balanus Hameri, lateral compartment, viewed externally. 

lb, „ „ carino-lateral compartment, viewed internally. 

Fig. 2a, Balanus bisulcatus, shell, 1^ nat. size. 

2b, ,, „ rostrum, viewed externally, to show the commencement of 

the folding of the walls, 1^ nat. size. 

2c, ,, „ var. plicatus, thrice nat. size. 

2d, ,, ,, very young shell, probably belonging to this species, enlarged 

about seven times. 

2e, 2f, „ „ scutum and tergum, external views. 

2g, 2h „ „ ,, ,, internal views. 

Fig. 3a, Balanus dolosus, rostrum, much enlarged, viewed internally. 

3b, ,, ,, portion of basal plate, much enlarged. 

3c, 3d, ,, ,, scutum and tergum, external views. 

Fig. 4<a, Balanus unguiformis, shell, twice nat. size. 

4Z>, „ „ shell, var. erisma, twice nat. size. 

4c, 4d, „ „ scutum and tergum, external views. 

4c, 4/, „ „ „ ,, internal views. 

Fig. oa, Balanus inclusus, shell, nearly thrice nat. size. 

5b, „ ,, basis of ditto, showing basal edges of the compartments. 

5c, ,, ,, var., with its rostro-carinal axis elongated. 

bd, „ „ var., „ „ „ showing the narrow 

furrowed basis and lower portions of the six compartments. 

5c, ,, ,, var. intermediate between the last two varieties, basal view of, 

nearly twice nat. size. 

5/, bg, „ ,, scutum and tergum, internal views from a very young 

specimen. 
Fig. 6a, Acasta undulata, rostrum viewed externally ; spinose var. ; twice nat. size. 

6(5, ,, ,, lateral compartment, smooth var., twice nat. size. 

6c, „ ,, basal cup. 

6d, „ ,, scutum viewed internally. 

6c, 6f„ „ ,, and tergum viewed externally. 

Fig. la, Pyrgoma Anglicum, shell, viewed from above, four times nat. size. 

lb, „ ,, scutum, internal view. 

7c, „ ,, tergum, internal view, engraved from recent specimen, this 

valve not having been found fossil. 



FOSSIL CFRRIPEDIA. 

Fig. 8a, Coroimla barbara, rostrum, external view ; nat. size. 

8(5, „ „ ,, viewed internally; twice nat. size. 

8c, „ ,, lateral compartment, internal view. 

Sd, ,, „ internal view, much enlarged, of small portion of basal margin 

of folded wall. 
Se, ,, „ external view, greatly enlarged, of small portion of surface of 

folded walls, near the basal margin ; (r r) the transverse 
ridges of growth. 
The following letters of reference apply to all the figures of Coronula : 

a, sheath marked trausversely in the upper part by the attachment of the 

opercular membrane. 
a', ala 

6, furrow on each side of (a), receiving the edge of the thick ala of the 
adjoining lateral compartment. 

c, special plate, on which the ala rests. 

d, radius, on the edge it may be just seen to consist of an outer layer (the 

normal radius), and a much thicker inner part (the pseudo or com- 
plementary radius) formed of oblique denticulated septa. 

e e' e", basal edge of wall, which from its commencement at e, or e", can be 
followed, folding up to near the basal edge of the sheath, to its 
termination at e" or e. 

ff, serrated lines of junction between the folds of the wall. 

Fig. 9a, Verruca Strom ia, much enlarged, engraved from a recent specimen, only certain 

valves having been found fossil. 
96, ,, ,, fixed scutum and tergum, internal views. The following 

letters apply to both these figures : 

a, rostrum. 

B, carina. 

s, moveable scutum, s', scutum fixed and modified so as to form part of shell. 

t, moveable tergum, t', tergum fixed, forming part of shell. 

In s, and s', a is the occludent margin ; b, the basal margin ; m, the plate 

to which the adductor muscle is fixed. 
In s and s', the tergal margin is marked by small dashes ; (') being the 
upper articular ridge, and (") the second or lower articular ridge : in 
s' (") is called the parietal portion of the valve. 
In t and t', the scutal margin is marked by small dashes ; (') being the first 
and upper articular ridge, hardly distinct from the occludent margin, 
and called in t' the occludent rim ; (") is the second or middle, and 
("') the lower or third articular ridge, called in t' the parietal portion of 
the valve : x is the carinal margin, called in t' the carinal rim, and z 
the basal margin. 
Fig. \9a, Verruca prisca, five times nat. size. 

\9h, 10c, ,, scutum and tergum, external views; engraved from a Belgian 

Cretacean specimen, the opercular valves not having been 
found in England. 






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